Terebratula Daleidensis - Royal Belgian Institute of Natural Sciences

BULLETIN DE L’INSTITUT ROYAL DES SCIENCES NATURELLES DE BELGIQUE
BULLETIN VAN HET KONINKLIJK BELGISCH INSTITUUT VOOR NATUURWETENSCHAPPEN
SCIENCES DE LA TERRE, 80: 47-84, 2010
AARDWETENSCHAPPEN, 80: 47-84, 2010
Re-examination of the Late Emsian rhynchonellid (brachiopod) Terebratula
Daleidensis RoemeR, 1844 from the Eifel area, and of some related species
by Paul SARTENAER
SaRtenaeR, P., 2010 – Re-examination of the Late Emsian
rhynchonellid (brachiopod) Terebratula Daleidensis RoemeR, 1844
from the Eifel area, and some related species. Bulletin de l’Institut
royal des Sciences naturelles de Belgique, Sciences de la Terre, 80:
47-84, 3 figs, 2 tables, 1 pl., Brussels, October 31, 2010 – ISSN
0374-6291.
Abstract
The Upper Emsian Terebratula Daleidensis RoemeR, 1844
(now Oligoptycherhynchus daleidensis) from the Daleiden
“Muldengruppe”, Eifel area, is scrutinized. The ubiquity and
considerable stratigraphic range gained by the species in the course
of time are exposed and questioned. After examination of the
lectotype, here designated, and rich existing collections, return to
the original definition of this abundant species is recommended.
Rhynchonella inaurita SandbeRgeR, G. & F., 1856, commonly
mistaken for daleidensis, is assigned to Inaequalibellirostrum
n. gen., the type genus of Inaequalibellirostridae n. fam. The
geographic distribution, the stratigraphic range, and affinities of
the type species of the new genus, I. pareti (de VeRneuil, 1850a)
from the middle part of the Emsian of the Cantabrian Cordillera
(Province of Leon) resulted, among others, in the reassessment of
the genus Stenorhynchia BRice, 1981, and its questionable presence
in the Armorican Massif.
Keywords: Oligoptycherhynchus daleidensis, Inaequalibellirostrum,
Inaequalibellirostridae, rhynchonellids, brachiopods,
Emsian, Eifel area, Cantabrian Cordillera, Armorican Massif.
Résumé
L’espèce Terebratula Daleidensis RoemeR, 1844 (aujourd’hui
Oligoptycherhynchus daleidensis) de l’Emsien Supérieur du groupe
de synclinaux (“Muldengruppe”) de Daleiden en Eifel est examinée
en détail. Son ubiquité et son extension stratigraphique considérable
acquises au cours du temps sont exposées et mises en doute. Après
examen du lectotype, ici désigné, et de riches collections existantes,
il est recommandé d’en revenir à la définition originelle de cette
abondante espèce. Rhynchonella inaurita SandbeRgeR, G. & F.,
1856, communément confondue avec daleidensis, est attribuée au
nouveau genre Inaequalibellirostrum, genre type de la nouvelle
famille Inaequalibellirostridae. La distribution géographique,
l’extension stratigraphique et les affinités de l’espèce type du
nouveau genre, I. pareti (de VeRneuil, 1850a) de la partie moyenne
de l’Emsien de la Cordillère Cantabrique (Province du Léon), sont
clarifiées. Deux résultats, parmi d’autres, en sont une réévaluation
du genre Stenorhynchia BRice, 1981 et la mise en question de la
présence de ce genre dans le Massif Armoricain.
Mots-clefs: Oligoptycherhynchus daleidensis, Inaequalibellirostrum,
Inaequalibellirostridae, Rhynchonellides, Brachiopodes,
Eifel, Cordillère Cantabrique, Massif Armoricain.
Introduction
Since its establishment as a characteristic species of
the famous Lower Devonian locality of Daleiden (Eifel
area), Terebratula Daleidensis RoemeR 1844 has
been abundantly quoted and discussed in the German
and world literature. Its considerable stratigraphic
range (Upper Ludlow to Strunian, chiefly Pragian and
Emsian) and its vast geographic distribution (Europe,
Asia, North Africa, Canada) indicate that the name
“daleidensis” has been unduly given to a long string of
forms. A critical scrutiny of the literature shows how
confusion progressively increased.
The aim of this paper is to advocate the return to
the original definition of the species (text, figures,
geographic and stratigraphic information) that was
clear and satisfactory for that time, and to restore its lost
identity and stratigraphic significance. The taxonomic
status of some species that have been mistaken for or
discussed in connection with daleidensis is examined
as well.

48 Paul SARTENAER
Major problems related in the past to daleidensis
Terebratula Daleidensis Roemer, 1844 versus T.
Daleidensis Schnur, 1853
A problem that popped up in the literature is the
difference between Terebratula Daleidensis described
by SchnuR [1853, p. 172, pl. 22, figs 1a-e (Remark: fig.
1e represents either another smaller specimen or is the
ventral view of the same specimen not reproduced to
scale)] and T. Daleidensis as originally established by
RoemeR (1844, p. 65, pl. I, figs 7a-c). Although some
authors, e.g. Dewalque (1868, p. 59; 1880, p. 67), and
RomanowSki (1880, p. 111), singled out a daleidensis
attributed to SchnuR. GoSSelet (1887, pp. 192-193,
194, 199) was the first to clearly state that one could,
pending a more complete study, separate SchnuR’s
4
Rhynchonella Daleidensis with median costae
3
5
from RoemeR’s R. Daleidensis with median costae.
GoSSelet went further in suggesting that SchnuR’s
R. Daleidensis could be Rh. inaurita SandbeRgeR, G.
& F., 1856 - he also used the expression Rh. inaurita
(daleidensis) SchnuR. He also considered that Rh.
Daleidensis RoemeR could be “synonyme, avec droit
de prénomination” of Rh. hexatoma SchnuR, 1851. In
this he was in opposition with SchnuR (1853), who
did not question RoemeR’s paternity of the species,
and so did most palaeontologists after GoSSelet, e.g.
Venyukov (1886, p. 108), and DReveRmann (1904,
p. 262). But the distinction survived for a long time,
e.g. Nalivkin (1930, pp. 61, 67), Paeckelmann &
SieveRtS (1932, pp. 53, 55), NöRing (1939, pp. 56,
57), and RzhonSnitSkaya (1973, pp. 31-33).
What were Gosselet’s arguments?
In his description of the species, RoemeR (1844, pl.
65, pl. I, figs 7a-c) mentioned the presence of 4 costae
on the fold, but figured a specimen showing 5 costae
on the fold on figure 7a, and 4 costae on figure 7b,
while 5 costae are drawn in the sulcus on the latter
figure. For his part SchnuR (1853, p. 172, pl. 22, figs
1a-e) considered the number of costae on the fold as
varying from 4 to 6 [5 and 6 being respectively the
number of costae in a rare variety, and in a stable
(“beständig”) smaller variety that could be a special
species (Remark: the small variety has been described
as Xahetomus hexadaleidensis by SaRtenaeR,
2009)], and illustrated a specimen carrying 4 costae
on the fold. Such contradictions, also underlined by
DReveRmann (1904, p. 262) and Maillieux (1931,
p. 23), have brought GoSSelet (1887, pp. 193, 194) to
4
conclude that SchnuR’s daleidensis with 4 costae on
the fold and 3 costae in the sulcus was different from
RoemeR’s daleidensis characterized by 5 and 4 costae,
respectively.
What about the specimen figured by Roemer (1844, pl.
1, figs 7a-c)?
In April 1980, the author had the privilege to examine
this specimen at the museum of the Institute of
Geological Sciences of the University of Wrocław,
where it is housed under the number 2054s, and in
August 2006 a cast (Pl. 1, Figs 6-10) was kindly sent to
him. This specimen matches RoemeR’s figures by size,
7 8
proportions, and number of lateral costae ( and ), and,
8
9
as such, is an acceptable representative of the species,
4
but it has median costae and wider costae than those
3
shown on the figures. This allows to cast doubts on its
reliability, although the label joined to it, and written by
RoemeR, indicates: “Devon. Daleiden in der Eifel. F.
Roemer leg. 1842.” Amongst the various explanations
put forward for explaining these contradictions, the
one proposed by DReveRmann (1904, p. 262), who
wrote that “die Abbildung RoemeRS...ist zweifellos
verzeichnet”, seems the most acceptable. As a matter
of fact, the large available collections of topotypical
material of daleidensis in old collections housed in
various museums, universities and scientific institutions
demonstrate that more than 75% of specimens show
4 costae on the fold as indicated by RoemeR himself
in his original description of the species, and 3 in the
5 4
sulcus. No single specimen shows or median cos-
5 5
tae. Costae have been counted on 172 specimens (see
Description of the species).
4
The conclusion is that is by far the usual number
3
of median costae in the material of the species described
by RoemeR (1844) as well as by SchnuR (1853), and
5
that GoSSelet’s (1887) estimate ( ) of this number in
RoemeR’s daleidensis is incorrect. However, GoSSelet
advocated other differences for separating SchnuR’s
daleidensis from RoemeR’s daleidensis. According
to him (pp. 192-193) the latter has a smaller size, a
very deep and pointed (“pointu”) sulcus, a trapezoidal
tongue, and generally no parietal costae (SchnuR’s
daleidensis show very rarely a parietal costa on each
side). With the exception of a very deep sulcus, these
differences are real, and SchnuR’s daleidensis should
be considered as a separate taxon for which, as already
suggested by GoSSelet (p.194), the name inaurita,
4

introduced by the SandbeRgeR brothers (1856, pp. 337-
338, pl. XXXIII, figs 5, 5a-c as Rhynchonella inaurita)
is available (see below). As far as parietal costae are
concerned, they have been often overlooked, and partly
rightly so, because the bounding costae of the fold and
the sulcus could be considered respectively as median
costae that are lower than the others, and lateral costae
that are higher than the others.
What about the localities of the specimens figured by
Roemer and Schnur?
We have seen that RoemeR’s figured specimen
(1844, pl. I, figs 7a-c) comes from Daleiden. This is
suggested anyhow by the name given to the species.
Furthermore, RoemeR (p. 65) mentioned the Daleiden
Grauwacke as his main collecting site, and added that
the species „ausserdem ist sie überall in der Grauwacke,
namentlich auch bei Waxweiler, Braubach, Coblenz,
Siegen, usw. verbreitet“ where it is „doch fast immer
sehr verdrückt“. Daleiden is cited by SchnuR only
as one of the localities where he found the species
(“Allenthalben in der Grauwacke, besonders häufig zu
Waxweiler, Daleiden, Prüm und Daun”); therefore it is
far from certain that his figures are those of one or two
(see remarks above) specimen(s) from this locality.
daleidensis = livonica and daleidensis versus livonica
von KeySeRling (1846, p. 240), BRonn (1848, pp.
1234, 1240-1241), and Kay S e R (1871a, pp. 314, 316,
list, p. 365; 1871b, pp. 518-520) considered Terebratula
Daleidensis identical to T. livonica von Buch, 1834
[Remark: after examination of Russian material,
Kay S e R (1889, p. 44) admitted that the two species
were different].
QuenStedt’s (1852, pp. 449-450, pl. 35, figs
42a-c; 1871, pp. 202-204, pl. 42, figs 57, 57a – 67 as
T. livonica) understanding of the species is broad and
obscure to say the least. Although the type locality of
T. livonica lies near Adsel, central Latvia, QuenStedt
(1852) described the species from “Dudley,
Grauwacken der Eifel, Gothland, etc.”, illustrated it by
a “Steinkern” from the “Grauwacke” of Daun (Eifel
area), and considered it “ganz vom Typus der Bicorner
des braunen Jura” [Remark: SaRtenaeR (1966, p. 3)
commented on the “Bicorner”].
QuenStedt (1871, figs 57, 57a) gave figures of a
specimen from the type area (“Mittellivland”), but, under
the heading T. Livonica, he discussed 13 species from
Daun, Dillenburg (Dill Syncline), Koněprusy (Bohemia),
and Gotland, and figured specimens from the first three
localities (figs 58-67). In particular, a “Böhmische”
Terebratula Daleidensis and related species
49
Livonica, different from the true (“ächte”) Livonica is
mentioned, and T. Daleidensis from Daun is considered
completely similar (“vollständig analog”) to T. livonica,
and not to be separated from it (“Ich trenne nicht”).
It is shown in the present paper that the daleidensis
from Daun is different from the type daleidensis of the
Daleiden “Muldengruppe” (Eifel area).
SteiningeR (1853, p. 58) thought that T. Daleidensis
was identical with the specimen of T. livonica figured
by de VeRneuil in MuRchiSon et al. (1845, pl. X,
figs 3a,b), but doubted that it was identical with T.
livonica as described by von Buch (1834), although
de VeRneuil’s specimen comes from the type locality,
Adsel in central Latvia.
The SandbeRgeR brothers (1856, p. 337, pl. XXXIII,
figs 5, 5a-c) incorporated in Rhynchonella inaurita not
only Terebratula Daleidensis and T. livonica, but also T.
Huotina de VeRneuil in MuRchiSon et al., 1845; after
examination of specimens of Rhynchonella livonica
from the Syas’ river, von SandbeRgeR (1889, p. 13)
sticked to the opinion that R. livonica was identical
to R. inaurita (= R. daleidensis). T. livonica expanded
still further when Kay S e R (1871b, pp. 518-520) and
Loewe (1913, pp. 63-65) attributed to it T. hexatoma,
Hemithyris Pareti de VeRneuil, 1850a and Terebratula
Wirtgeni SchnuR, 1853 in addition to the three species
already included in it by the SandbeRgeR brothers;
Kay S e R stated that Rhynchonella livonica “gehört zu
den an meisten veränderlichen Rhynchonellen” and
added that the “pugnaceenförmig hexatoma mit 5
scharfen Falten im Sinus...ist nichts Anderes als eine
kleine Fortläuferin der Daleidensis [Grauwacke bei
Daun und Daleiden] im Kalke”.
Venyukov (1886, pp. 108-109, 122-123) added
R. turanica RomanowSki, 1880 to the list of species
assimilated to R. livonica by Kay S e R.
BaRRoiS (1882, p. 267) expressed a finer opinion
when he declared R. daleidensis closely related (“très
voisine”) to R. livonica, R. inaurita, and R. Pareti,
although, forty years later, BaRRoiS et al. in GoSSelet
et al. (1922, p. 97) included R. nympha, R. daleidensis,
R. livonica, R. Pareti, R. sub-Pareti, and R. sublivonica
in a group “composé de formes intimement
alliées, sinon identiques entre elles” [Remark: BaRRoiS
et al. considered QuenStedt (1871) the author of
sub-livonica and referred to his figures (pl. 42, figs
67, 68). QuenStedt mentioned only Terebratula
pseudolivonica, which is a species of BaRRande
(1847); fig. 67 is, according to him, a “Zwischenform”
between pseudolivonica and Livonica, that cannot be
separated with certainty, and fig. 68 is T. cuneata from
Gotland]. Therefore, sub-livonica is a misquotation.

50 Paul SARTENAER
MauReR (1896, p. 657) was less accommodating
in expressing his conviction that the “forms” from the
Rhenish Lower Devonian could not be considered as
being the same species as R. livonica. Focusing on R.
daleidensis and R. livonica, GüRich (1909, p. 146)
wrote: “Da diese Arten im Unter- und Mitteldevon im
Fluss sind, hält es schwer Grenzformen als besondere
Arten heraus zu greifen”. In adopting a wider vision,
Reed (1922, p. 95), who had already (1921, p. 317)
recognized “a considerable range of variation” in R.
daleidensis, considered that R. daleidensis, R. inaurita,
R. Pareti, R. Partridgiae WhidboRne, 1897 and
R. turanica “should be regarded as synonyms of R.
livonica, or at any rate not worthy of ranking above
varieties”. This was the last attempt to mistake various
species for the Lower Frasnian species Ripidiorhynchus
livonicus, as the Latvian species is known nowadays;
these species, of which some are discussed in the
present paper, are independent taxa.
inaurita versus daleidensis
When the SandbeRgeR brothers (1856, pp. 337-338,
pl. XXXIII, figs 5, 5a-c) established Rhynchonella
inaurita, they included in it three previously described
species (Terebratula livonica, T. Daleidensis, and
T. Huotina), and, thus, the priority should have been
given to the oldest known species, T. livonica. At the
same time, they gave to the new species a considerable
stratigaphic range - Oriskany Sandstone (Pragian) to
Stringocephalenkalk - and a vast geographic distribution:
France (Boulonnais, Normandy), Germany (Dill
Syncline, Eifel, Lahn Syncline, middle Rhine valley,
Taunus), Latvia, Pensylvania, and Russia (Central and
Main Devonian Fields, Pechora, W Siberia). After
examination of specimens of Rhynchonella livonica
from the Syas’ river, von SandbeRgeR (1889, p. 13)
dropped any doubt he had so far concerning the identity
of R. livonica with R. inaurita (= R. daleidensis), and
stated that he believed, like Venyukov (1886), that
there was no palpable (“greifbare”) difference between
the two species.
Rhynchonella inaurita has usually been considered
a junior synonym of Terebratula Daleidensis, e.g. by
KRantz (1857, p. 150), RomanowSki (1880, p. 111),
GoSSelet (1887, p. 194), MauReR (1896, p. 656),
DReveRmann (1904, p. 262), GüRich (1909, pl. 45,
figs 6a-d), who went as far as choosing SandbeRgeRs’
figures (pl. XXXIII, figs 5, 5a-c) of Rhynchonella
inaurita to illustrate R. daleidensis, and Maillieux
(1931, p. 24). DavidSon (1870, pp. 72, 75, 78-81,
87, pl. 5, figs 1-3; 1881, p. 341, pl. 38, fig. 21?, figs
35a,b) recognized the species in the Upper Devonian
of Devonshire. This strange and false occurrence
was apparently based on identification by GoSSelet.
Kay S e R (1871b), Venyukov (1886), and Loewe (1913)
assigned the species to R. livonica (see above). Kay S e R
(1878, pp. 142-143) recognized its validity, but later
(1889, pp. 44-45) questioned separation of R. inaurita
and R. daleidensis. BaRRoiS (1882, p. 267) considered
R. inaurita as “très voisine” of R. daleidensis, R.
livonica, and R. Pareti, and Reed (1922, p. 95) took
the position indicated above under the discussion of
daleidensis versus livonica.
As already indicated, the author considers inaurita
as a valid taxon, and in so doing, he falls in with
GoSSelet’s (1887, p. 194) suggestion that the name
inaurita could be given to Terebratula Daleidensis
as described by SchnuR (1853) if it is accepted that
it is different (“si on admet cette différence”) from T.
Daleidensis as originally described by RoemeR (1844)
(see above). Of course, one will have to forget the
synonymy, the stratigraphic range, and the geographic
distribution proposed by the SandbeRgeR brothers for
the species they established.
The species is included in a new genus in the present
paper.
daleidensis versus pareti
When establishing Hemithyris Pareti, de VeRneuil
(1850a, pp. 160, 162, 177) mentioned that the species
was distinct from Terebratula daleidensis only «par la
dépression qu’elle présente des deux côtés du crochet».
de VeRneuil (1850b, table, pp. 780-781, p. 785; 1866,
p. 11) upheld this opinion, and added, in 1866, that there
would be «peu d’inconvénients à (les) confondre». In
the meantime SchnuR (1853, p. 172) agreed that the
two species were close to each other (Remark: SchnuR
wrote mistakenly Pailleti instead of Pareti).
As already mentioned, Kay S e R (1871b, pp. 518,
520), Venyukov (1886, pp. 108, 123), and Loewe
(1913) incorporated Hemithyris Pareti in T. livonica,
and Kay S e R (1878, p. 143) confirmed that the two
species were “täuschend ähnlich”. BaRRoiS (1882,
p. 267; 1889, p. 85) also considered Rhynchonella
Daleidensis, R. inaurita, R. livonica, and R. Pareti as
“très voisines” [Remark: BaRRoiS et al. in GoSSelet
et al. (1922, p. 97) went further in including R. nympha,
R. daleidensis, R. livonica, R. Pareti, R. sub-Pareti,
and R. sub-livonica in a same group (see above)].
OehleRt (1884, pp. 415-416) believed that R. Pareti,
R. subpareti, OehleRt 1884, and R. cypris d’ORbigny,
1847 «forment un groupe naturel avec Rh. livonica,

Rh. nympha, Rh. pseudolivonica, etc., et ne sont sans
doute qu’un même type, modifié dans le temps et dans
l’espace». Reed’s (1922, p. 96) stand on the subject
has been reminded above. Kay S e R (1889, p. 45) while
maintaining that R. Pareti was “nicht verschieden”
from R. livonica, added that the former species
“schliesst sich der grossen Abänderung von Daleiden
[R. daleidensis]...an”; DReveRmann (1904, p. 262)
and HüffneR (1917, p. 312) declared themselves in
full agreement with Kay S e R. From then on R. Pareti
has consistently been introduced into the synonymy of
Camarotoechia daleidensis, e.g. by Maillieux (1931,
pp. 20-21, 24; Maillieux’s synonymy is a list of
citations and not a critical synonymy), Comte (1938,
pp. 58-59, 86), Renaud (1942, pp. 98-99), Le MaîtRe
(1944, p. 48), LlopíS Lladó (1961, pp. 265-267), and
Schumann (1965, p. 93).
Things changed when palaeontologists started to
work actively in the Cantabrian Cordillera, from where
the species was first recorded, and where it is recognized
as a valid taxon with a known stratigraphic range.
The species is designated in the present paper as the
type species of a new genus.
daleidensis versus hexatoma
This problem is more important than the preceding
ones, because it still persists in modern literature and
hampers progress in classification and stratigraphy.
We have seen that Kay S e R (1871b), Venyukov
(1886), and Loewe (1913) incorporated T. hexatoma
with T. livonica. Kay S e R (1871b, p. 520) considered
T. hexatoma as a “Fortläuferin” of T. Daleidensis, from
which it takes over, and reaffirmed (1889, p. 43) his
opinion in writing: “Die [Rhynchonella daleidensis]
am Rhein sehr verbreitete, durch das ganze Unterdevon
hindurchgehende und auch in das Mitteldevon der Eifel
(hexatoma SchnuR) aufsteigende Art”. GoSSelet
(1887, p. 194) went further in suggesting that R.
hexatoma and R. Daleidensis (not R. Daleidensis
as described by SchnuR 1853; see above) could be
synonyms, but a few years later (in USSheR, 1890, p. 498;
1903, p. 27) he expressed himself slightly differently
in writing that R. daleidensis was near (“voisine”) to
R. hexatoma. This was also ViëtoR’s (1918, p. 439)
position, who considered R. daleidensis as a “nahe
verwandte Form” of R. hexatoma. In the meantime
GüRich (1896, p. 284) made out of Terebratula
hexatoma a variety of RoemeR’s “Hauptform” (T.
daleidensis), but MauReR (1896, p. 659) explicitly
wrote that there were no transition forms between R.
daleidensis and R. hexatoma. DReveRmann (1904, p.
Terebratula Daleidensis and related species
51
262) and FuchS (in SpRieSteRSbach & FuchS, 1909,
pp. 69-70) put an end to these wavering views in stating
that Rhynchonella hexatoma was a “besondere Art” or
“selbständige Art” as originally established by SchnuR
(1851). This should have closed the controversy, but
Schmidt (1941) and Le MaîtRe (1952a, b) reopened
the debate.
Schmidt (1941, pp. 7-11) called “daleidensis-
Gruppe” or “Gruppe der Camarotoechia daleidensis” a
group of forms giving the impression of unity (“Eindruck
der Einheitlichkeit”), although she recognized that
shape was a very variable character (“die Gestalt ist
innerhalb weiter Grenzen sehr wandelbar”). She focused
her attention on the Lower and Middle Devonian
representatives of the group, which, according to
her, was distributed throughout the whole Devonian.
Schmidt stated that C. hexatoma includes a number
of variations (“Abweichungen”), which she treated as
subgroups (“Untergruppen”) or subspecies that showed
transitions (“Übergänge”) between them. With this in
mind she described three subspecies (C. hexatoma
hexatoma, C. hexatoma soetenica Schmidt, 1941, and
C. hexatoma wetteldorfensis Schmidt, 1941) based on
the number of costae and the value of the apical angle
as distinctive characters. At the same time, she showed
that the subspecies were not clear-cut; a particular
specimen could often not be attributed to one or another
of them, and that exceptional specimens were not
rare. More generally, she stressed that the subgroups
of the “daleidensis-Gruppe” had to be considered as
provisional.
On top of the restrictions and approximations
contained in Schmidt’s scheme, the cumulated
stratigraphic range of the subspecies, i.e. of the species,
is considerable and therefore suspicious. These are
an uppermost Emsian (Heisdorf Formation in the
Prüm and Hillesheim Synclines) and, questionably,
lowermost Eifelian (Lauch Formation in the Prüm
Syncline) for hexatoma wetteldorfensis; lower part
of lower Eifelian [“Geeser Horizont” of the Nohn
Formation in the Hillesheim Syncline, not to mention
a “non-characteristic specimen from the “Geeser
Horizont” (upper part of the middle Eifelian Ahrdorf
Formation) in the Gerolstein Syncline], and middle
Eifelian “Sötenicher Schiefer” in the Sötenich Syncline
(according to Schmidt, 1936) for hexatoma hexatoma;
and (lower) (Lauch and Nohn formations) and middle
Eifelian (Ahrdorf and Junkerberg Formations) to upper
Givetian in the Blankenheim and Sötenich Synclines
for hexatoma soetenica.
The specimen of hexatoma wetteldorfensis figured
by Schmidt (1942, figs 3a-c, p. 393), and duplicated by

52 Paul SARTENAER
WeRneR (1980, fig. 16, p. 17), is completely different
from the holotype figured by her in 1941 (pl. 1, figs
2a,b).
The Upper Emsian species Rhynchonella imitatrix
FuchS in SpRieSteRSbach & FuchS, 1909 from the
Bergisches Land and R. (Wilsonia) dillensis FuchS,
1914 from the Dill Syncline are not identical to
hexatoma wetteldorfensis, a possibility set forth by
Schmidt (1941, p. 9); according to her, R. imitatrix
could be a juvenile of hexatoma wetteldorfensis. These
two remarks indicate that Schmidt’s conception of the
subspecies was unclear.
According to the author, the subspecies hexatoma
soetenica, that covers most of this range, contains
more than one taxon, a possibility already envisaged by
Schmidt (p. 11) for the specimens from the Sötenich
Syncline. Furthermore, although the subspecies is
present in two synclines, all figured and sectioned
specimens come from the Givetian of a restricted area
(Sötenich and surroundings) in the Sötenich Syncline:
holotype (pl. 1, figs 4a-c) and one paratype (pl. 1,
figs 5a, b) from the “Stringocephalen-Kalk” and two
paratypes (pl. 4, figs 57a, b; pl. 5, figs 5-8) from the
“Stringocephalen-Schichten”.
As concerns hexatoma hexatoma, SaRtenaeR
(2007, p. 45) reminded that the “Geeser Horizont” near
Üxheim in the Hillesheim Syncline is of early Eifelian
age (lower “Nohner Schichten”), and therefore not
equivalent, contrary to Schmidt’s statement, to the
“Geeser Horizont” of the Gerolstein Syncline, which
is located in the upper part of the Ahrdorf Formation
(middle Eifelian).
After examination of the original collections
the author concludes that the external characters of
the subspecies hexatoma soetenica and hexatoma
wetteldorfensis differ considerably from those of
hexatoma hexatoma, although some internal features
show some analogy. They belong to two different
genera, none of them being Oligoptycherhynchus
SaRtenaeR, 1970, the genus to which hexatomus and
daleidensis belong. Is Cupularostrum SaRtenaeR,
1961 the genus to which Camarotoechia hexatoma
soetenica should be assigned? LaRdeux & MoRzadec
(1979, p. 23 as “Camarotoechia” soetenica) elevated
the subspecies to the rank of a species that BRice
(in BRice & MoRzadec, 1983, pp. 549-550, 553-
555, pl.1, figs 1a, b, 2-7) described and attributed to
Cupularostrum. This is an acceptable proposition
for the Lower Givetian (Tibidy Formation) 40 (~)
specimens from two sections near Le Faou (“Rade
de Brest”, “Département du Finistère”, Armorican
Massif), but it does not apply to the original Eifelian
material, which is difficult to tackle. The holotype
(XVII 323c) is stratigraphically (“Stringocephalen-
Kalk”) and geographically (Sötenich) unsatisfactorily
located. The largest collection of hexatoma soetenica
comes from the middle Eifelian Ahrdorf and Junkerberg
Formations in the central part of the Sötenich Syncline;
it is particularly abundant in the Dalbenden horizon
(third from base of the four recognized in the Junkerberg
Formation) (PauluS, 1961a, pp. 424, 427-429, 432;
1961b, pp. 32, 38; Dickfeld, 1968, unpublished thesis:
“Stratigraphie und Fauna im Westteil der Sötenicher
Mulde”). Dalbenden is only two kilometres to the east
of Sötenich. It is surprising that Schmidt (1941, p.
11) mentioned as coming from the “Stringocephalen-
Schichten” an old and large collection she examined in
the “Landesmuseum” (Berlin), because the path Urft-
Gïrzenberg from where this collection derives is also
in Dalbenden or close to it. It is also remarkable that
similar material coming from the same formation has
been collected from the near-by Blankenheim (OchS
& WolfaRt, 1961, pp. 23, 28, 30) and Rohr (GlinSki,
1961, pp. 279, 281, 283) Synclines. In short, a thorough
examination is needed for clarifying the stratigraphic
position and range of the subspecies in its type area,
and for dealing with the various taxa that have been
identified as such in France (“Rade de Brest”; Lower
Givetian), Germany (Sauerland; Middle Eifelian),
in Libya (Fezzan; Couvinian and Givetian-Frasnian
transition beds), in Mauritania [Adrar; Givetian-
Frasnian boundary beds (probably Frasnian)], and
in Spain (Cantabrian Cordillera; Efelian-Givetian
transition beds).
Le MaîtRe (1952a, p. 112) stated that C. daleidensis
presented “variations”. The same year (1952b, p. 328)
she called “groupe de C. daleidensis” “toute une série
d’espèces” gathered around the species in the whole
Devonian of Europe and North America. According
to her, in Europe the Lower Devonian section (C.
daleidensis and C. nympha) of this group was replaced
in the Middle and Upper Devonian by “toute une série
de formes voisines” showing “variations” in various
characters: C. elliptica, C. hexatoma, C. hexatoma
soetenica, C. hexatoma wetteldorfensis, C. triloba
fornicata, C. ferquensis, C. letiensis, C. omaliusi, C.
triaequalis. Le MaîtRe admitted presence of similar
species (“espèces similaires”) in the United States
of America in the Lower Devonian (C. oriskania),
and the Middle and Upper Devonian (C. congregata,
C. congregata var. parkheadensis, C. horsfordi,
C. orbicularis, and C. prolifica). She (Le MaîtRe,
1952b, p. 329) also recognised in Mauritania (Adrar)
three “types principaux” or “groupes”, one of them

attributed to C. hexatoma, in mentioning that such a
specific assignment was difficult on account of the
“modifications qui survienent chez une même espèce”.
With Schmidt (1941) and Le MaîtRe (1952a,b)
confusion reached a peak, and the “groupings” they
proposed be better forgotten. Unfortunately confusion
survived.
The multiplication of subspecies, varieties,
variations, and transition forms cannot be regarded as an
advance in our knowledge, but rather as the wavering of
our understanding in the absence of a strong support.
In order to unravel this imbroglio, we have to take
up the matter anew. This is what DRot (1964, pp.
182-187, figs 76-78, pl. 18, figs 8a-c, pl.19, figs 10a,b,
11a-c; 1981, pp. 46-48, figs 2a,b, p. 47, pl. 1, figs 3a-c,
4a-c, 5a-c, 6a-c) thought when she stumbled against
the problem of the alleged presence of Camarotoechia
daleidensis and C. hexatoma in the Morrocan pre-
Sahara. She declared (1981) that the “groupe de
formes” including these two species and C. hexatoma
wetteldorfensis was in need of a “révision générale”.
Hexatoma (subspecies and varieties included) is
mentioned in the literature with a Pragian to Lower
Famennian stratigraphic range from the following
countries: Germany (Bergisches Land, Dill Syncline,
Eifel area, Harz Mountains, Hörre, Lahn Syncline,
Mosel valley, middle Rhine valley, Sauerland, Taunus),
Belgium, Canada, England, France, Iran, Libya,
Mauritania, Morocco, Poland, Russia, Spain, and
Usbekistan. Eifelian to Lower Frasnian and Upper
Emsian to Lower Eifelian are the given ranges for
hexatoma soetenica and hexatoma wetteldorfensis,
respectively.
Most of identifications and ranges, if not all,
are incorrect, with, of course, the exception of the
stratum typicum and locus typicus, the Gees horizon
(“Geeser Horizont”) (middle Ahrdorf Formation, lower
Middle Eifelian), and the Gees-Pelm area (Gerolstein
Syncline). The species, which is very rare, is also found
in the lower Nohn Beds (upper Lower Eifelian) of the
Hillesheim Syncline.
The Couvinian material (eleven “crushed”
specimens, “most of them very flat”) from
Grzegorzowice (northern Holy Cross Mountains)
identified as Camarotoechia hexatoma by Bi e R n at
(1954, p. 489, table 1, pp. 490-491, pp. 509-511, pl. IV,
figs 6-10) shows some analogy to the Eifelian species.
It differs from it in its small thickness, and especially in
its septalium partly covered with two thin plates with
bifurcated extremities, a structure never encountered
by the author in any other rhynchonellid.
Terebratula Daleidensis and related species
Varieties, mutations and subspecies of daleidensis
53
Varieties (not formally named)
Many varieties of daleidensis have been mentioned in
the literature, although seldom formally named.
The two varieties of Terebratula Daleidensis from the
“Grauwacke” singled out by SchnuR (1853, p. 172)
have been mentioned above. One of them has been
described as Xahetomus hexadaleidensis by SaRtenaeR
(2009, pp. 35-37, pl.1, figs 11-60).
Other varieties, mutations, etc. have been proposed
by various authors:
- variety [FuchS, 1899, pp. 57, 78, from the Lower
Emsian of the Lorelei area (Middle Rhine valley);
1911, p. 713 from the Daadener Schichten (Lower
Emsian) of the Siegerland; in SpRieSteRSbach
& FuchS, 1909, pp. 6, 70, from the Upper Emsian
(“Remscheider Schichten”) of Bergisches Land;
DahmeR, 1921, table, p. 198, table, p. 300, from
the Upper Emsian (“Remscheider Schichten” of
Bergisches Land and “Rammelsberger Schichten”
and “Nessigi-Schichten” of Harz Mountains; 1932,
pp. 373, 383 from the Middle Siegenian of Juseret
(Neufchâteau Syncline), 1942, p. 268, fig. 29, p.
287, from the Lower Emsian of Ziegenberg (eastern
Taunus)];
- variety? (DReveRmann, 1904, p. 262, from the
Siegenian of Seifen (Westerwald);
- varieties [RomanowSki, 1880, p. 111, from the
Devonian of Turkestan (Chatkal river valley);
GüRich, 1909, p. 146, from the Middle Devonian;
Maillieux,1936, pp.14, 87 quoting DahmeR (1932),
and p. 88, two varieties not worth being described
(“trop mal représentées pour être utilement décrites”)
from the Upper Siegenian of Fauvillers (Neufchâteau
Syncline)];
- mutations (FuchS in SpRieSteRSbach & FuchS
1909, p. 70, from the Upper Emsian of the Eifel and
middle Rhine valley areas);
- “Abänderungen” [Kay S e R, 1889, pp. 44-45, from
the Lower (“Unter-“) and Upper (“Ober-“) Emsian
(“- coblenzschichten”) of Eifel area (Daun, Daleiden),
includes GoSSelet’s (1887) suggestion to separate
SchnuR’s Terebratula Daleidensis from RoemeR’s
T. Daleidensis (see above); DahmeR, 1921, p. 208,
from the Upper Emsian (“Schalker Schichten”) of
Harz Mountains];
- “Abart” [Paeckelmann & SieveRtS, 1932, p. 55,
from Paphlagonia (Asia Minor); DahmeR, 1921, p.
278, from the Lower Emsian of the middle Rhine
valley (Lorelei area) and the Upper Emsian of Harz
Mountains; 1932, p. 383, from the Middle Siegenian

54 Paul SARTENAER
of Juseret (Neufchâteau Syncline); 1942, p. 270, from
the Lower Emsian of Ziegenberg (eastern Taunus)];
- “Zwischenstufe” between RoemeR’s (1844) T.
Daleidensis and SchnuR’s (1853) T. hexatoma
[GüRich, 1896, p. 284 and Sobolev, 1909, p. 508,
from the “Spiriferensandstein” of the northern Holy
Cross Mountains (Miejska Góra)];
- small variety [Kay S e R, 1895, p. 208, pl. III, figs 1-2
from the Lower Couvinian of Pepinster, considered
as a juvenile specimen of Rhynchonella daleidensis;
Dewalque (in de PieRpont, 1895, pp. 169, 171, 173,
174), and de DoRlodot (1901, p. 174, foot-note 1, p.
182, p. 182, foot-note 1), from the Lower Couvinian
of the Vesdre Massif (Pepinster) and the southern
flank of Lustin (Hestroy, Burnot) and Godinne
(Rouillon) Anticlines in the Meuse river valley in the
central part of the Dinant Basin].
Varieties (formally named)
The variety Rhynchonella daleidensis gracilior from
the Lower Emsian of the Lorelei area (middle Rhine
valley) was proposed by FuchS (1899, pp. 69, 74, 84,
86, 88) and considered by him as “sehr bezeichnend für
die höheren Niveaus der Hercyniaezone [“Singhofen-
Schichten]”. The variety has also been mentioned: 1)
from the Upper Emsian of the Bergisches Land by
fuchS in SpRieSteRSbach & FuchS [1909, pp. 6, 70
as a variety similar (“derartig”) to the variety gracilior;
one crushed specimen], 2) from the Upper Emsian
(“Kahlebergsandstein”, Giengelberger Schichten”,
“Nessigi-Schichten”, “Schalker Schichten”, “Festenburger
Schichten”, and “Rammelsberger Schichten”of
the Harz Mountains by DahmeR [1921, table, p. 174
possibly, p. 191, table, p. 198, pp. 206, 277-279, pl.
15, figs 18-20 (= three specimens from the “Nessigi-
Schichten”), table, p. 300; 1946, p. 179]. This
identification was confirmed by FuchS in DahmeR,
1921, who compared it with the small Belgian variety
of daleidensis from the Lower Couvinian of Pepinster
described by Kay S e R (1895, p. 208). FuchS, however,
referred to Kay S e R’s pl. III, figs 1-4, that Kay S e R
considered as juvenile (pl. III, figs 1-2) and “normal”
(pl. III, figs 3,4) forms of daleidensis (Remark:
ASSelbeRghS, 1923, p. 25, pl. I, figs 4-9, attributed the
specimens identified as Camarotoechia daleidensis by
Kay S e R to C. imitatrix); 3) from the Lower Couvinian
on the southern border (Olloy area) of the Dinant Basin
by Maillieux (1939, p. 2 as C. gracilior); and 4) from
the Emsian of Paphlagonia (Asia Minor) by HeRitSch
& von GaeRtneR (1929, pp. 191-193, pl. 1, figs 12-
13), and Paeckelmann & SieveRtS (1932, p. 55).
Maillieux (1931, pp. 23, 25 as C. gracilior)
contemplated the possibility that Camarotoechia
imitatrix and Rhynchonella daleidensis gracilior could
be synonyms.
MittmeyeR (2008, table 2, p. 142, pp. 171, 173 as
Oligoptycherhynchus daleidensis gracilior) has given
a fresh impetus to the variety in mentioning it in the
middle Lower Emsian Singhofen Substage (i.e. in the
lower part of the Lower Bendorf Beds), the lower part
of the Spitznack Beds (lower part of the Singhofen
Substage = stratum typicum of the variety), and in the
Wambach Beds (new lithostratigraphic unit). These
beds extend, according to MittmeyeR, to western
Westerwald and the Mosel river region, the Taunus
and eastern Hunsrück, and the southern Taunus,
respectively. MittmeyeR (2008, p. 203) considers O.
prodaleidensis n. sp. a forerunner (“Vorläufer”) of O.
daleidensis gracilior (see below).
FuchS’ (1899) gracilior is a nomen nudum; it has
neither been described nor defined in accordance
with Article 12 of the ICZN. This is also the case for
DahmeR’s (1921) gracilior, because expressions such
as “eine kleine zierliche Form von der Verwandtschaft
daleidensis” (Kahlebergsandstein, Upper Emsian,
Harz Mountains) or “eine zwerghafte Abart” (Lower
Emsian, Lorelei area, middle Rhine valley) cannot be
considered as descriptions or definitions; specimens of
figures 18-20, plate 15, are also not an indication (Article
12c). DahmeR himself admitted (p. 278) that a closer
description (“eine nähere Beschreibung”) of the variety
did not exist. The name gracilior may be made available
later with a new authorship. DahmeR submitted his
collection from the Harz Mountains to FuchS, who
found it very much alike (“übereinstimmend”) gracilior
from the middle Rhine valley. Such conformity needs
to be substantiated. Furthermore all small forms with
4
median costae cannot be put in the same basket no
3
matter their age (early Emsian, late Emsian, and early
Couvinian) and their geographic location (middle Rhine
valley, Taunus, Hunsrück, Westerwald, Bergisches
Land, Harz Mountains, and Dinant Basin).
For the time being, the relation of gracilior
with the late Emsian daleidensis from the Daleiden
“Muldengruppe” (Eifel area), and its assignment to
the middle Eifelian genus Oligoptycherhynchus is not
acceptable. MittmeyeR intended to give a description
and an illustration of gracilior based on his large
collections from the Singhofen Substage (stratum
typicum), but could not do this in the context of the
compendium on the Devonian of Germany, published
in 2008.
The subspecies O. daleidensis minor (FuchS)

mentioned by MittmeyeR (2008, p. 190, from the
Lower Bendorf Beds is a misprint for O. daleidensis
gracilior.
Prodaleidensis Mittmeyer, 2008
Oligoptycherhynchus prodaleidensis was mentioned by
MittmeyeR (1997, p. 22) as a nomen nudum from the
Reudelsterz Beds (upper part of the Ulmen Substage)
south of Schutz near Manderscheid in the Eifel area
south of the “Eifelkalkmulden”, and considered as a
forerunner (“Vorläufer”) of O. daleidensis. MittmeyeR
(2008, table 2, p. 142, pp. 163, 169, 186-190, 203, pl.
1, figs 17-18) described the species, a forerunner of
O. daleidensis gracilior, from the same area and from
Taunus and Hunsrück, with the following stratigraphic
range: lower (Kaltenborn Beds) and upper (Ramersbach
Beds) parts of the Middle Siegenian; lower (Nitztal
Beds) and uppermost (Saxler Beds) parts of the
Upper Siegenian; lower (Eckfeld and Sauerthal Beds)
and upper (Lower and Upper Reudelsterz, and Kaub
Beds) parts of the Lower Emsian. The description and
illustration (internal moulds of one brachial valve and
one ventral valve) of the species are unsatisfactory,
possibly due to the publication in a compendium. It
is hoped that the species will be better described and
figured in the future with proper comparison with
known taxa. At this stage it can be stated that the
only connection of the species with daleidensis is the
4
number ( ) of median costae, and that its assignment
3
to the genus Oligoptycherhynchus is erroneous.
Unnamed species and subspecies
Solle (1976, table, p. 22) mentioned Oligoptycherhynchus
n. sp. aff. daleidensis from the lower
“Gladbach-Schichten” of the lower part of the
“Vallendar-Schichten” (= upper Lower Emsian) of the
Olkenbach Syncline.
FuchS (1982a, fig. 10, p. 253, p. 254 as “large growth
forms of Oligoptycherhynchus n. sp. aff. daleidensis”;
in FuchS & PluSquellec 1982, p. 26 as “une nouvelle
espèce voisine d’ O. daleidensis”) proposed a new
species, never described, from the Singhofen Substage
(middle Lower Emsian) and the lower and middle
Vallendar Substage (upper Lower Emsian) of the Eifel
area east of the “Eifelkalkmulden”; in the same area
and south of the “Eifelkalkmulden” FuchS (1989,
pp. 111, 115) declared O. daleidensis “bezeichnend”
for the Ulmen Substage (lower Lower Emsian), and
singled out a “small form” from the Nasingen Beds
(lower beds of the Ulmen Substage) and a “grosse
Variante” (O. sp. aff. daleidensis) from the Neuerburg
Terebratula Daleidensis and related species
55
Beds (upper beds of the Ulmen Substage). These
various forms will have to find a taxonomic position,
distinct from daleidensis as described in the present
paper, if located in FuchS’ collection housed in the
“Senckenberg Forschungsinstitut und Natur Museum,
Frankfurt am Main”.
WenndoRf (2001, p. 20) mentioned O. daleidensis
subsp.? from the “Emsquarzit” (base of the lower Upper
Emsian Lahnstein Substage).
Number of median costae of daleidensis used as a
stratigraphic tool
RoemeR (1844) and SchnuR (1851), the founders of
Terebratula Daleidensis and T. hexatoma, respectively,
considered the number of dorsal median costae, four
for the former species, and six for the latter, as a distinct
character. Since then, this costal numbers played a
predominant part in identifying, grouping or separating
species, subspecies and varieties showing evident or
alleged similarities to these two species.
Two authors in particular, MauReR (1896) and
Schmidt (1941) attributed a stratigraphic significance
to the number of median costae.
MauReR (1896, pp. 656-659) stressed the point
that Rhynchonella daleidensis was one of the most
abundant species of the Lower Devonian east of the
Rhine that showed extraordinary equal contour and
size, and limited “Formschwankungen” at every level.
He stated that specimens from the lower “Stufen”
(“untere Grauwacke”, “Haliseritenschiefer”, and
4
“Coblenzquarzit”) had median costae (with the
3
exception of one specimen from the quartzite with
5
median costae) and those from the upper “Stufen”
4
(“Chondritenschiefer”, “Hohenrheiner Stufe”, and
6
“Cultrijugatus-Stufe”) (with the exception of three
5
5
specimens from the “Cultrijugatus-Stufe” with and
4
4 median costae).
3
MauReR reminded that GoSSelet (1887) envisaged
the possibility of separating SchnuR’s (1853)
4
Terebratula Daleidensis with median costae from
3
5
RoemeR’s T. Daleidensis with median costae
[Remark: GoSSelet suggested in that case that the
name inaurita proposed by the SandbeRgeR brothers
(1856) could apply to the former]. In this context,
according to MauReR, Rhynchonella inaurita and R.
daleidensis would characterize the predominant form
of the lower part and the upper part of the Lower
4

56 Paul SARTENAER
Devonian, respectively. Such an interpretation, in
contradiction with the
6
median costae advocated for the
5
upper “Stufen”, is not a fair assessment of GoSSelet’s
opinion (see above), because at that time by most
palaeontologists the two species were considered of
the same age (“Grauwacke”, “Spiriferensandstein”),
and identical.
In her discussion of the Camarotoechia daleidensis
group, ranging from the late Emsian (Wiltz Beds)
to the late Givetian, Schmidt (1941, pp. 8, 10)
4
separated specimens with median costae from the late
3
Emsian (Wiltz Beds and lower part of the Wetteldorf
6
Sandstone) from specimens with median costae
5
from the latest Emsian (Heisdorf Beds), Eifelian, and
Givetian beds [Remark: the mention by Schmidt of
6
specimens with median costae in the Heisdorf Beds
5
is due to the presence of a subspecies of C. hexatoma,
C. hexatoma wetteldorfensis, that consequently allows
to give a early to middle Devonian age to the species;
see above]. Exceptions to this scheme are explained by
Schmidt in the following way: isolated (“vereinzelte”)
6
specimens from the Wiltz Beds with median costae
5
could come from a very high horizon in these beds
(Remark: these specimens are included in the species
Xahetomus hexadaleidensis), and herald the “6-costae
clothing” (“6-Rippen-Tracht”) of higher beds; the
Upper Eifelian or Lower Givetian specimens of C.
4
hexatoma soetenica that have median costae from
3
near Blankenheim are otherwise nevertheless in such
conformity (“Übereinstimmung”) with the “6-rippigen
Sötenicher Form” that “eine Abtrennung nicht möglich
ist, zumal da die Gesamtzahl der Rippen die gleiche
ist”.
These are only two examples of the damaging
consequences resulting from focusing essentially on
one external character (number of median costae),
and neglecting all others. The picture becomes blurred
and judgement is affected. An extreme example, from
which any stratigraphic consideration is excluded,
is the following one: Maillieux (1931, pp. 21-24)
stated that BaRRoiS’s (1889, pp. 86-87, pl. V, figs 2a-e)
Rhynchonella nympha from the Armorican Massif,
5
having median costae could just be (“il est possible
4
qu’elle doive être rapportée à”) Rhynchonella Pareti,
4
a species with median costae that he includes in the
3
synonymy of Camarotoechia daleidensis. Therefore
5
it is only on account of the median costae that
4
Maillieux questionably included BaRRoiS’s taxa,
possibly identical according to him, in the synonymy
of Camarotoechia daleidensis, implying that this
4
restriction would be lifted if these taxa had median
3
costae.
As for daleidensis, the two following remarks must
4 6
be borne in mind. Firstly, although and represent
3 5
the number of median costae of a large majority of
specimens of daleidensis and hexatoma, respectively,
other ratios exist and indicate some degree of variability.
Secondly, there are more rhynchonellid species (and
4 6
genera) with and median costae than just dalei-
3 5
densis and hexatoma known from the Emsian and
Eifelian. Within the Wiltz Beds (Upper Emsian) alone,
there is one species of which the majority of specimens
4
possesses median costae (daleidensis), and another
3
6
of which the majority of specimens possesses median
5
costae (Xahetomus hexadaleidensis = one of the two
varieties of daleidensis mentioned by SchnuR, 1853,
p. 172).
Stratigraphic range and geographic distribution of
daleidensis according to the literature
Daleidensis has been commonly mentioned in beds
ranging in age from the Siegenian to the early Eifelian;
it has also been occasionally mentioned in the Lower
Ludlow, Gedinnian, Upper Devonian, Frasnian, and
Strunian.
Rhenish facies E and W of the middle Rhine valley
In Bergisches Land, Dill Syncline, Hunsrück,
Kellerwald, Lahn Syncline, Mosel valley, Siegerland,
Taunus, and Westerwald, daleidensis has been reported
from the whole Lower Devonian (e.g. Kay S e R, 1889,
p. 43, table, p. 110, 1908, pp. 139, 143; MauReR,
1896, p. 657; ViëtoR, 1918, p. 438, table, p. 466)
or from the middle and upper Lower Devonian (e.g.
Paeckelmann & SieveRtS, 1932, p. 55). It means:
Lower and Upper Siegenian, and Lower and Upper
Emsian (Hunsrück), Lower and Upper Siegenian,
and Lower Emsian (Siegerland), Middle Siegenian
(Westerwald), Upper Siegenian and Lower Emsian
(Kellerwald, Taunus), Upper Siegenian and Lower
and Upper Emsian (Mosel valley), Lower and Upper
Emsian (Dill Syncline), Lower and Upper Emsian, and
Lower Eifelian (Bergisches Land, Lahn Syncline).

Eifel area
- In the Eifel area considered as a whole, daleidensis
has been mentioned from the “Cultrijugatusstufe” by
Schulz (1883, table, p. 167), from the lower Middle
Devonian by FuchS in SpRieSteRSbach & FuchS
(1909, p. 80), and from the Upper Emsian Beds by
SpRieSteRSbach (1942, table, p. 90).
- In the “Eifelkalkmulden”, aside of the Daleiden
“Muldengruppe” (type area, see below under the
description of daleidensis), the species has been
mentioned from the Lower Devonian of the Sötenich
Syncline by QuiRing (1915, pp. 92, 107, table
between p. 160 and p. 161), from the Upper Emsian
Wetteldorf Beds of the Gerolstein Syncline by Heibel
(1969, table 2, p. 548), from the Upper Emsian of the
northern margin of the Prüm Syncline by weRneR
(1969, table 2, pp. 170-171, table 3, pp. 180-182,
184, table 4, p. 189, p. 218), and from the Lower
Emsian and Upper Emsian Wetteldorf Sandstone of
the Prüm Syncline by GoSSelet in Duponchelle
(1880, pp. 326-327), ASSelbeRghS (1913, p. 156;
1928, pp. 9-10) and various authors (e.g. Schmidt,
1942, p. 390, tables 1, 2, p. 401; Bultynck 1970,
pl. XXXVII), respectively. Kay S e R (1871b, p. 520;
1889, p. 43, table, p. 110) accepted the presence
of the species also in the Middle Devonian of
the “Eifelkalkmulden”, because (see above) he
considered the Eifelian Terebratula hexatoma as the
“Fortläuferin” of T. Daleidensis.
- Region SE and E of the “Eifelkalkmulden”. The
presence of daleidensis has been mentioned in the
Middle Siegenian (MittmeyeR, 1982, pp. 30, 33, 35,
36, 39-41), in the Middle and Upper Siegenian, and
Lower Emsian (SimpSon, 1940, p. 11, 18-22, table
4, p. 60, table 8, p. 66-67), in the Upper Siegenian
and the three substages (Ulmen, Singhofen, and
Vallendar) of the Lower Emsian [e.g. Maillieux,
1910, table, p. 207, p. 218; RödeR, 1960, pp. 10, 11,
18, 20, 22-24, 29, 30, 32, 33, 35-38, 40, 41, 43, 44,
46, table 1, p. 50, pp. 53, 54, 57; FuchS (1974, pp.
25, 32, 33, table 1, pp. 35-36, p. 40, table 2, pp. 44,
46, pp. 50, 56-59, 61, 62, 64, 68, 71, 74, 77, 78, 82,
83, 85, 86, 88, 95, 97-100; 1982a, pp. 233-237, fig.
10, p. 253, p. 254; 1982b, pp. 157, 159, 161, 164,
169, 172; Negendank, 1983, pp. 21-24, 27, 29-30,
149; KölSchbach et al., 1993, table 1, pp. 306-
308], in the Lower Emsian (e.g. FRech, 1889, p. 195;
DReveRmann, 1902, p. 103; DienSt, 1914, table,
pp. 604-605; Mauz, 1935, pp. 23, 78; Solle, 1956,
pp. 8, 10, 11; RödeR, 1957, pp. 134- 135, 137, 138;
negendank, 1974, fig. 9, pp. 22, 24, 89; Dohm,
1976, fig. 9, p. 19; KowalSki, 1976, pp. 236-237,
Terebratula Daleidensis and related species
57
241-244; FüRSich & HuRSt, 1980, p. 307; FuchS
& PluSquellec, 1982, p. 26; FuchS, 1989, p.111;
MittmeyeR, 1997, pp. 22-23), in the Upper Emsian
(“Stufe der Spirifer cultrijugatus”) (e.g. Schulz,
1883, table, p. 167), in the Lower and Upper Emsian
(e.g. Solle, 1937, pp.7, 9, 11, table, p. 17, p. 22,
table, p. 34, table, p. 43, table 1; 1976, table, p. 22,
pp. 43-44, table, pp. 52-53, table, pp. 90-91, table, p.
109, p. 117, table, p. 125, table, p. 139).
- Region W and SW of the “Eifelkalkmulden”.
Daleidensis has been reported from the middle
Emsian of Üttfeld and Brandscheid areas, and the
Upper Emsian of Irrhausen by ASSelbeRghS (1928,
pp. 9, 15), the Nasinger Schichten (lowermost Lower
Emsian) and the Stadtfelder Schichten (uppermost
Lower Emsian) by NöRing (1939, pp. 69-72), and
the three subdivisions (from base to top: “Nasingen-
“, “Niederraden-“, and “Neuerburg-Schichten”) of
the lower Lower Emsian Ulmen Substage, of which
daleidensis is characteristic (“bezeichnend”), and the
two subdivisions (“Altscheuern-“ and “Merlbach-
Schichten”) of the middle Lower Emsian Singhofen
Substage of the Neuerburg area by FuchS (1989, p.
111-117, 119-122).
- Region N of the “Eifelkalkmulden”. Daleidensis has
been mentioned by SaueR (1968, pp. 500-501) in the
Lower Emsian of the neighbourhood of Zülpich.
Regions close to the type area
Not too far away from the type area, daleidensis has
been reported from the Upper Emsian and Lower
Couvinian of the Vesdre Massif and from the Lower
Siegenian to the Lower Emsian of the Aachen area. The
species has been mentioned from the Middle Siegenian
to the Upper Emsian of the Oesling Basin (Grand
Duchy of Luxemburg), and from the Lower Siegenian
to the Lower Couvinian of the Ardenne [Dinant Basin
(Belgium and France), Ardenne Anticline, Neufchâteau
Syncline, Laroche-Houffalize Syncline, Bastogne
Anticline, and eastern border of the Stavelot Massif
(Belgium), and northern border of the Givonne Massif
(France)]; a single Frasnian occurrence on the southern
border of the Dinant Basin is due to MouRlon (1881,
p. 15). In Sauerland, the species is known from the
Upper Emsian, and in the Harz Mountains from the
Lower and Upper Emsian, and the Lower Eifelian.
Other regions of the world
According to the literature, the stratigraphic range
and the geographic distribution of daleidensis is as
follows: Algeria (Siegenian, Lower and Upper Emsian,

58 Paul SARTENAER
Strunian), Canada (New Scotland, Pragian), China
(Inner Mongolia, Upper Emsian), England (Siegenian
to Eifelian), France (Gedinnian, Siegenian, Emsian,
Lower Couvinian), Iran (Siegenian to Upper Emsian),
Kazakhstan (Emsian, Eifelian), Libya (Lower Devonian,
Givetian-Lower Frasnian), Mauritania (Siegenian
to Lower Couvinian), Morocco (Lower and Upper
Siegenian, Lower and Upper Emsian, Lower Eifelian),
Poland (Upper Ludlow, Upper Emsian), Rumania
(Gedinnian, Siegenian), Spain (Gedinnian, Siegenian,
Emsian), Turkestan (Upper Devonian), Turkey [Lower
Devonian (Siegenian, Emsian)], Uzbekistan (Upper
Devonian).
Conclusions
Before trying to get out of the fix we are in, it is useful
to examine clearly the succession of steps that allowed
the confusion to progressively install itself in a process
that is common for many species of old standing:
- wrong understanding, and thus incorrect
identifications (often based on one or two external
characters) that resulted in incorporating of various
forms in the Eifel species Terebratula daleidensis
and T. hexatoma, and in proposing arbitrary
groupings; the word “group” being used with various
meanings;
- erroneous and not substantiated generic assignments;
- spreading of the two species from their loci typici
(Daleiden “Muldengruppe”, and Gerolstein Syncline,
respectively) to other parts of the Eifel area, to
other regions of Germany (both species: Bergisches
Land, Dill Syncline, Hörre, Harz, Lahn Syncline,
Moselle valley, middle Rhine valley, Sauerland,
Taunus; daleidensis alone: Aachen area, Hessisches
Hinterland, Hunsrück, Kellerwald, Siegerland,
Westerwald), to bordering and near-by regions
(both species: Ardennes, Dinant Basin, France,
Grand Duchy of Luxemburg; daleidensis alone:
Vesdre Massif; hexatoma alone: Namur Basin), to
other European countries (both species: England,
Poland, Russia, Spain; daleidensis alone: Rumania;
hexatoma alone: Kuznetsk Basin), and to the rest
of the world (both species: Iran, Libya, Mauritania,
Morocco, Uzbekistan; daleidensis alone: Algeria,
China, Kazakhstan, Nova Scotia, Turkey ; hexatoma
alone: Canadian Arctic);
- considerable stratigraphic range: for daleidensis:
Eifel area [Siegenian to Lower Eifelian (chiefly
upper part of Lower Emsian, and Upper Emsian)],
other regions of Germany [Siegenian to Lower
Eifelian (chiefly Lower and Upper Emsian)], and
other parts of the world [Gedinnian to Middle
Eifelian, and Frasnian (chiefly Siegenian to Emsian)]
not to mention exceptional occurrences in beds of
late Ludlow, Gedinnian, Frasnian, and Strunian
ages; for hexatoma: Eifel area [Lower Emsian to
Givetian (chiefly Upper Emsian to Middle Eifelian)],
other regions of Germany (Upper Emsian to Lower
Givetian), and other parts of the world (Pragian to
Lower Givetian, Frasnian, and Lower Famennian);
- quasi general lack of any information on the precise
location or range of species within lithostratigraphic
units, rarely smaller than a formation, with, as a
result, the impossibility to trace eventual transition
forms and to follow evolutionary lineages;
- usual preservation as moulds of the fossils obtained
from the Rhenish “Grauwacke” involving more
disadvantages than advantages.
Systematic Palaeontology
Family Sapphicorhynchidae SaRtenaeR, 2007
Genus Oligoptycherhynchus
SaRtenaeR, 1970, emend. 2007
SaRtenaeR (2007, p. 45) acknowledged that “the
wide range (middle Siegenian to Eifelian) advocated
by him in 1970 for Oligoptycherhynchus has become
unacceptable”, with, accordingly, the elimination from
the genus of all other species than the type species,
the Middle Eifelian O. hexatomus. The revaluation
of the Upper Emsian daleidensis in the present paper
convinced the author that this species had to remain
in the genus, and that its assignment to a new genus
suggested by him (SaRtenaeR, 2009, p. 34) had to be
refuted.
Type species: Terebratula hexatoma SchnuR, 1851.
Diagnosis
The emended diagnosis of the genus, which is here
assigned to he family Sapphicorhynchidae, is as
follows: medium size; outline subcordiform; uniplicate;
inequivalve; top of dorsal valve at front; commissure
sharp; well marked sulcus and fold, starting at a short
distance from the beaks; sulcus moderately deep with
flat to slightly convex bottom, wide at front; high fold
with slightly convex top; high subrectangular tongue
with sharp borders, sometimes recurved posteriorly
in its upper part; ventral beak slightly to strongly

incurved; narrow ventral interarea; costae in moderate
number, regular, simple, moderately elevated to
elevated, angular or angular with rounded top, starting
at the beaks; parietal costae common; apical angle
wide; moderately thick shell; moderately thick dental
plates, diverging in the apical region, becoming parallel
to convergent anteriorly, long and moderately thick
septum; deep septalium, amphora-shaped in transverse
serial sections, covered by a moderately robust
connectivum; hinge plate composed of two horizontal
to slightly concave parts; inner ridges of dental sockets
high; teeth short and robust.
Oligoptycherhynchus daleidensis (RoemeR, 1844)
Pl. 1, Figs 1-35
Figured specimens are deposited in the collections of the
Royal Belgian Institute of Natural Sciences, Brussels,
with registration numbers prefixed IRScNBa.
Lectotype, locus typicus, stratum typicum
The only syntype figured by RoemeR (1844, pl.
I, figs 7a-c) from the “Grauwacke” near Daleiden
(“Daleiden Muldengruppe”, Eifel area) is here selected
as the lectotype of the species, and reproduced (Pl.
1, Figs 6-10). Four specimens identified in 1843 by
RoemeR are deposited in the “Hauptsammlung” of the
“Paläontologisches Museum, Museum für Naturkunde,
Berlin”,
Other material
The 665 (~) specimens examined come from the
stratum typicum and the locus typicus: 500 (~)
specimens in the collections of the “Preussische
Königlich geologischen Landesanstalt” housed in the
”Paläontologisches Museum, Museum für Naturkunde,
Berlin” (nine of them = topotypes A-I, IRScNBa12703-
12711); 31 specimens deposited in the Department of
Palaeontology of the British Museum (Natural History),
London, purchased or accepted as a gift in the beginning
of the twentieth century; 141 specimens being part of
the collections of the “Senckenberg Forschungsinstitut
und Natur Museum “, 35 of them collected by J. Schnur
(the specimen XVII 180w figured by Schmidt, 1941,
pl. 1, figs 1a-c, belongs to this collection), and 99 by K.
Jaeger. All specimens are internal moulds.
Description
Remarks: On account of the following three
conclusions reached in the present paper no description
of Oligoptycherhynchus daleidensis after 1844 is
reliable, although the descriptions may include
Terebratula Daleidensis and related species
59
some pertinent features: 1) the difference between
Rhynchonella inaurita and Terebratula Daleidensis
sensu SchnuR, 1853, a possibility already envisaged
by GoSSelet (1887) (see above), resulting in the
assignment of Rhynchonella inaurita to a new genus,
Inaequalibellirostrum, described below; 2) the absence
of the species in Germany outside the Eifel area; 3)
the absence of the species in other countries of the
world. Therefore, our understanding of the species
relies on the original description by RoemeR (1844, p.
65, pl. I, figs 7a-c), which contains some of the most
characteristic features of the species. RoemeR’s figures
are also the best ones existing so far, in spite of the slight
discrepancy in the number of median costae mentioned
in the text and seen on one of the three figures (fig. 7b);
this problem has been discussed above. Photographs
(Pl. 1) of six specimens and the plaster cast of the
lectotype of O. daleidensis show that the species was
originally well figured and well described.
None of the many specimens examined by the
author showed a preserved shell, although according
to RoemeR (1844, p. 65), the species was represented
by “zahlreichen völlig unverdrückten Exemplare,
zum Theil mit erhaltener Schale”. This is the reason
why there are no published transverse serial sections
[Remark: Schumann (1965, fig. 22, p. 95), made
sections from a specimen of Rhynchonella pareti
(now Inaequalibellirostrum pareti) mistaken for
Camarotoechia daleidensis]. Nevertheless, most of
the internal features of the species can be inferred
from moulds; the author confirmed the presence of a
connectivum in making sections in a specimen.
What follows refers only to specific characters in
need of further elaboration. Width of sulcus at front
between 54 and 73%, mostly between 57 and 67%, of
shell width. Maximum shell width located between 52
and 72%, mostly between 62 and 67%, of shell length
anterior to ventral beak. Apical angle between 112°
and 126°, mostly 112° to 118°. Ratios
l
: 0.70 to 0.90,
w
t
mostly 0.70 to 0.82; : 0.80 to 0.97, mostly 0.85 to
w
t
0.97; : 1 to 1.31, mostly 1 to 1.16. The general costal
l
formula in median, parietal, and lateral categories
4
derived from at least 75% of the specimens is: ; 0;
3
7 - 9
3
. The ratios of costae are given in Table 1. Width
8 - 10
of median costae near front between 2.5 and 3 mm.
Measurements of ten specimens, of which seven
have been photographed, are given on Table 2.
Columns 3, 5-7 refer to specimens of usual size. The
distorted specimen figured by SchnuR (1853, pl. 22,

60 Paul SARTENAER
Number
of costae
Table 1 – Number of median and lateral costae of Oligoptycherhynchus daleidensis (RoemeR, 1844).
figs 1a-d) falls into a small category of particularly
wide specimens such as topotype I, IRScNBa12711
(column 10). The enlarged (x1.5) specimen figured by
Schmidt (1941, pl. 1, figs 1a-c) is similar to the one of
figures 31-35 of Plate 1.
The percentage of specimens from the type area
5
with median costae includes those that have been
4
1 2
3
2
4
3
5
4
5 6
qualified as rare by SchnuR (1853, p. 172), and by
FuchS in SpRieSteRSbach & FuchS (1909, p. 70), who
considered them as mutations. SteiningeR (1853, p.
58) mentioned one specimen with five, another with six
costae on the fold, and even one with six costae in the
sulcus. Schmidt (1941, p. 8) mentioned “vereinzelte
Stücke in den Wiltzer Schichten haben 6 Rippen auf
dem Sattel”; these specimens belong to the species
Xahetomus hexadaleidensis.
Comparisons
The major difference between the abundant Upper
Emsian O. daleidensis and the rare upper Lower and
lower Middle Eifelian O. hexatomus is the number of
4
6
median and parietal costae ( ; 0 for O. daleidensis; ;
3
5
1–1
for O. hexatomus); the number of lateral costae is
1–1
7–9
similar ( ) [Remark: the total number of costae, 16
8–10
Median costae Lateral costae
Number of
specimens
12
10
313
20
5
360
%
3.3
2.8
87
5.5
1.4
100
Number
of costae
5
6
7 6
7
8
8
9
9
10
10
11
11
12
12
13
Number of
specimens
1
7
64
107
103
28
6
1
317
%
0.3
2.2
20
33.8
32.5
8.9
2
0.3
100
to 18, indicated by Schmidt (1941, p. 8) for hexatomus
is a lapsus calami for 26 to 28]. Parietal costae are very
rare in O. daleidensis, but the external costae of fold are
commonly slightly lower than the others, and the ventral
internal lateral costae slightly higher than the others.
In O. hexatomus, ventral flanks flatten or even reverse
their curvature near the antero-lateral commissures,
where they end as spurs, the sulcus is generally slightly
deeper, the anterior commissure is lower than the
front margin, and the frontal parts of sulcus and fold
form together a convex surface with flattened costae.
The largest specimens of O. hexatomus never reach
the size of the largest specimens of O. daleidensis.
Schmidt (1941, pl. 5, figs 4a-i as Camarotoechia
hexatoma hexatoma) published transverse serial
sections from one specimen of hexatoma coming
from the lower Nohn Beds (upper Lower Eifelian)
of Üxheim (Hillesheim Syncline). More detailed
sections (including the crura) from another specimen
(IRScNBa12712) from the same locality are given in Fig.
1. As already mentioned, the “Geeser Horizont” of the
Nohn Beds is older than the “Geeser Horizont” of Gees
and Pelm (Gerolstein Syncline), which is the stratum
typicum and the locus typicus of Oligoptycherhynchus
hexatomus. But, because no topotype of the species
has been spotted so far in existing collections, the

in mm
l
lvv unrolled
w
t
tvv
tdv
l/w
t/w
t/l
Apical angle
Topotype
G
IRScNBa
12709
13
24
17.9
14.8
5.2
9.6
0.73
0.83
1.14
117°
Topotype
A
IRScNBa
12703
14.9
30
18.8
18.1
5.5
12.6
0.79
0.96
1.20
116°
Topotype
B
IRScNBa
12704
17.3
34
19.2
18.6
6
12.6
0.90
0.97
1.08
112°
Topotype
D
IRScNBa
12706
18
36.5
20.7
20.9
5.7
15.2
0.87
1
1.16
114°
15.7
Terebratula Daleidensis and related species
Table 2 – Measurements of ten specimens of Oligoptycherhynchus daleidensis (RoemeR, 1844). Abbreviations: l = length; w =
width; t = thickness; vv = ventral valve; dv = dorsal valve.
author, like Schmidt, had to be contented with making
sections from a rare specimen presented to the author
by Dr. K.-W. Wenndorf (collected by Berd Trost)
that is externally similar to the specimen figured
by SchnuR (1853, pl. 23, figs 2a-e as Terebratula
hexatoma). This specimen is here formally designated
as the lectotype of the species. It could be argued that
such a designation already took place. As a matter of
fact, Schmidt (1941, p. 9) declared that among the
subspecies of Camarotoechia hexatoma described by
her, C. hexatoma hexatoma seemed to be the typical
one, as best corresponded to SchnuR’s figures (“scheint
mir die typische zu sein, denn sie entspricht am besten
der Abbildung bei SchnuR”). Such a statement cannot
be considered as the designation of a lectotype, because
it does not comply with Article 74.7 of the ICZN.
Moreover, the German word “Abbildung” used in the
singular covers the two syntypes figured by SchnuR
(figs 2a-e and figs 2f, g, plate 23). With the designation
of a lectotype, the specimen of figures 2f, g becomes a
paralectotype. This status is questionable in the present
case, because SchnuR, as explained by SaRtenaeR
(2009, p. 35), figured this specimen under the name
Terebratula hexatoma, but described it (SchnuR, 1853,
p. 172) as one of the two varieties of T. Daleidensis.
Anyhow, paralectotype or not, this specimen has been
chosen as the holotype of Xahetomus hexadaleidensis.
The Lower Givetian species Sapphicorhynchus
sappho (Hall, 1860) and Oligoptycherhynchus
daleidensis share several characters, e.g. a strongly
dorsibiconvex profile; the dorsal valve thickest
posterior to front; deeply serrated antero-lateral and
lateral commissures; a short interarea; well marked
sulcus and fold starting at a short distance from the
Topotype
C
IRScNBa
12705
30
20.8
17.6
5.5
12.1
0.75
0.85
1.12
121°
Lectotype
18.2
31
21.1
18.3
5
13.3
0.86
0.87
1
115°
Topotype
H
IRScNBa
12710
17.7
31
22.3
17.9
5.2
12.7
0.79
0.80
1.01
113°
Topotype
E
IRScNBa
12707
17.3
36.5
23.9
22.8
6.2
16.5
0.72
0.95
1.31
118°
Topotype
F
IRScNBa
12708
20.1
37
24.6
21.9
5.4
16.5
0.82
0.89
1.09
118°
18.6
37.5
26.4
23
5
18
0.70
0.87
1.24
126°
61
Topotype
I
IRScNBa
12711
beaks; a wide, moderately deep sulcus, with flat to
slightly convex bottom; a clearly delineated tongue
slightly recurving posteriorly in its uppermost part; the
top of fold gently convex; a moderate number of well
marked, wide, simple costae starting from the beaks;
similar internal characters (detached dental plates, long
septum, deep and wide septalium, undivided hinge
plate, connectivum).
Many characters make Sapphicorhynchus sappho
distinct from Oligoptycherhynchus daleidensis: a
t t t
smaller thickness indicated in the
w
and ratios ( :
l w
0.63 to 0.80, mostly 0.66 to 0.80, for S. sappho against
0.79 to 0.99, mostly 0.79 to 0.93, for daleidensis);
t
:
0.75 to 1, mostly 0.88 to 1, for sappho against 0.95
to 1.12, mostly 0.95 to 1, for daleidensis); a more
constant outline; a greater width*; the maximum shell
width located more posteriorly (45 to 65% of shell
length anterior to ventral beak for sappho against 60
to 65% for daleidensis); the ventral flanks flattening
near the front*; the anterior commissure fused into
the wall formed by the extremities of costae; a lower
tongue with transverse-subtrapezoidal outline; the
top of tongue lower than maximum shell thickness; a
5–6
different general costal formula ( 5 − 6 ; 0 to 11–0
− 0 and
4–5 4 − 5 11–0
− 0
1–1 1−
1 6–8
; 6 8
4
7 – 9
; 0; for daleiden-
1–1 1−
1
− for sappho against
77–9
− 9
3
8 –10
sis); the common presence of parietal costae; a wider
angle of the cardinal commissure; more delicate internal
features (the preservation as moulds of the specimens
of daleidensis does not allow a detailed examination of
most features, and, in particular, of the crura). (* = only
valid for middle-large specimens of sappho).
l

62 Paul SARTENAER
Fig. 1 –
Oligoptycherhynchus hexatomus (SchnuR, 1851). Camera lucida drawings of serial transverse sections; figures are
distances in mm forward of the ventral umbo. IRScNBa12712. Üxheim (Hillesheim Syncline, lower Nohn Beds, upper
Lower Eifelian). Measurements: length = 16.3 mm; width = 21.7 mm; thickness = 15.8 mm.
Trigonirhynchia CoopeR, 1942 differs from
Oligoptycherhynchus in most characters. The genus is
only mentioned here, because Schmidt (1965, p. 6),
although underlining obvious differences, considered
that Camarotoechia daleidensis and C. hexatoma had
some features (outline, profile, costae) in common with
its type species Trigonirhynchia fallaciosa Bay l e,
1878 (Pragian, Armorican Massif).
Stratigraphic position and geographic location
In a remote past, Oligoptycherhynchus daleidensis
has been mentioned in the type area (Daleiden
“Muldengruppe” and adjoining region of the Grand
Duchy of Luxembourg = type area stricto sensu, and
Prümer Syncline = type area lato sensu) from the
“Spiriferensandstein”, the Daleiden-, Eifel-, Hierges-,
Rhine (Rhenish)-, and Waxweiler-“Grauwacke”, and
from the Daleiden-Waxweiler beds. A more precise
lithostratigraphic unit, covering the upper part of the
“Grauwacke”, the “Oberk(c)oblenzschichten” came
progressively in use.
O. daleidensis is now restricted to the Upper
Emsian Wiltz Beds (“Wiltz-Schichten”), of which it is
a characteristic species (see e.g. StRuve, 1964, p. 227;
WeRneR, 1980, p. 14; KowalSki, 1983, p. 101).
The presence of the species in Germany outside
the type area has still to be substantiated. The undue
importance given to a given number of median costae
4
), considered the most important character, if not the
(
3
only one, of O. daleidensis, has led to the assignment
to that species of a long string of Siegenian and
Emsian forms. No specimen has been spotted in the
various collections examined by the author. Evidence

is lacking for the abundant isolated and embedded
valves from the “Singhofen-Gruppe” (middle Lower
Emsian) of the Taunus (near the “Loch-Mühle” NW
Gemünden) assigned by KutScheR & MittmeyeR
(1970, pp. 43-44, 45-46, 49, pl. 15, figs 11-14) to the
“Normalform” of Daleiden, to which they remarkably
(“in bemerkenswerter Weise”) correspond, for the only
4
reason that they show an almost constant number ( )
of median costae.
3
The presence and the stratigraphic range of
daleidensis in other parts of the world (see above) must
be disregarded.
Inaequalibellirostridae n. fam.
Type genus: Inaequalibellirostrum n. gen.
Diagnosis
Shell of medium to medium-large size, with right angle
triangle profile, subtrigonal outline, and convexoconcave
ventral valve; apical angle moderately wide;
thickest at front margin; top of shell = top of tongue;
well marked lunulae; short ventral interarea; pointed,
projecting ventral beak; sulcus, fold, and costae start
from beaks; sulcus wide and deep; fold high; tongue
high; costae well marked, in moderate number,
sharp, high, simple, regular, deeply indenting the
commissure; no parietal costae; dental plates separated
from the wall by wide umbonal cavities; long septum;
wide, deep, cupula-shaped septalium, covered by
a thin connectivum in its anterior part; undivided
hinge plate; raduliform crura; high internal socket
ridges; well marked denticula; and slighty impressed,
longitudinally elliptical and narrow ventral muscle
field.
Comparisons
The new family Inaequalibellirostridae differs from the
family Sapphicorhynchidae in a subtrigonal outline;
a right angle triangle profile; a convexo-concave
ventral valve; better marked lunulae; a pointed,
projecting ventral beak; a deeper sulcus; a higher fold
with strongly convex top; sharper, higher costae; the
absence of parietal costae; a larger, wider, and deeper
septalium (connectivum occupies more than half the
width of hinge plate); teeth entering the dental sockets
vertically in serial transverse sections (not laterally
as in the family Sapphicorhynchidae); dental plates
divergent posteriorly, subparallel anteriorly, i.e. not
convergent; higher internal socket ridges; and inverted
L-shaped crura in their distal part. [Remark: constancy
Terebratula Daleidensis and related species
63
of the last two differences needs confirmation; other
internal characters shown in Fig. 3 are supported
by the serial transverse sections and drawings to
be found in the literature (DRot, 1964, fig. 75, p.
181 as “Camarotoechia” pareti; Schumann, 1965,
fig. 22, p. 95 as C. daleidensis, standing for pareti,
and WeStbRoek, 1967, fig. 7, p. 8, fig. 32, p. 30 as
Trigonirhynchia pareti)].
Before comparing the families Inaequalibellirostridae
and Trigonirhynchiidae Schmidt, 1965, some
points need to be clarified.
Reasons for re-examining the contents of the
family Trigonirhynchiidae, and its five subfamilies
(Hemitoechiinae Sava g e, 1996, Ripidiorhynchinae
Sava g e, 1996, Rostricellulinae Rozman, 1969,
Trigonirhynchiinae Schmidt, 1965, and Virginiatiinae
AmSden, 1974) have been examined at length by
SaRtenaeR (2007, pp. 42-43, fig. 1, p. 48).
A first step in that direction has been the elevation
by SaRtenaeR (2001, p. 208; 2003, p. 183) of
the subfamily Ripidiorhynchinae to the family
rank, with Ripidiorhynchus SaRtenaeR, 1966 as
sole representative. The genera Cyphoterorhyncus
SaRtenaeR, 1964, Hemiplethorhynchus von Peetz,
1898, and Pseudosinotectirostrum Yudina, 1991
are excluded from the family Ripidiorhynchidae.
At the same time, SaRtenaeR chose not to include
in the family various genera resulting from a
revision of that genus, and to establish monogeneric
families. Therefore, the following genera were
provisionally included, with reservation, in the
family Trigonirhynchiidae: Hypselororhynchus
SaRtenaeR, 2001, Kedridorhynchus SaRtenaeR,
2001, Orophomesorhynchus SaRtenaeR, 2001,
Piridiorhynchus SaRtenaeR, 2001, Poleomesorhynchus
SaRtenaeR, 2001, Paropamisorhynchus
SaRtenaeR, 2001, Porthmorhynchus SaRtenaeR,
2001, Saxuli-rostrum SaRtenaeR, 2001, and
Gesoriacorostrum, SaRtenaeR, 2003.
Sava g e (2007, pp. 2703-2707) added to the four
genera included earlier by him (Sava g e, 1996, table
3, p. 256; 2002, pp. 1074-1077) in the subfamily
Ripidiorhynchinae the eight genera proposed by
SaRtenaeR (2001) plus Hunanotoechia Ma, 1993,
while the genus Gesoriacorostrum was omitted. This
is acceptable for some of these genera as long as they
are not included in a subfamily (Ripidiorhynchinae)
of the family Trigonirhynchidae, but in the family
Ripidiorhynchidae as recommended by SaRtenaeR
(2001, 2003).
On the other hand, Sava g e suggested that
Hypselororhynchus and Kedridorhynchus were

64 Paul SARTENAER
synonyms of Porthmorhynchus and Saxulirostrum
respectively. Diagnoses of these genera by SaRtenaeR
(2001, pp. 199-201, 203) indicate that differences
between these genera are important.
The Middle-Upper Frasnian Porthmorhynchus
ferquensis (GoSSelet, 1887) from Boulonnais can
easily be separated from Hypselororhynchus farsani
(BRice in BRice & FaRSan, 1977) (Middle-Upper
Frasnian, western Afghanistan) by a large size;
stronger and higher costae; a usually smaller number
of median and lateral costae; parietal costae only
occasionally present; clearly detached dental plates
and septum; and wide umbonal cavities. The Upper
Frasnian Saxulirostrum saxatilis (Hall, 1867) from
NC Iowa can easily be separated from Kedridorhynchus
cedarensis (StainbRook, 1942) (Upper Givetian, NE
and CE Iowa) by a larger size; a deeper sulcus; higher
fold, tongue, and costae; slender internal structure; and
an undivided hinge plate.
These are only two of the 43 genera synonymized
by Sava g e (2002, 2007). With the exception of the few
genera already considered synonyms by specialists,
most of these synonymies are not accepted by the
present author. Neither are most of the 52 nomina
dubia of Sava g e in the same publications [SaRtenaeR
(2004, p. 8) commented on Platyglossariorhynhus
considered a nomen dubium by Sava g e]. On the
other hand, some genera considered for a long time
as synonyms have been treated as valid genera. Thus,
Bergalaria Schmidt, 1975, considered a synonym
of Flabellulirostrum SaRtenaeR, 1971 by DRot
(1982, p. 74), BRice in BRice & MoRzadec (1983,
pp. 549, 563), and SaRtenaeR (1985, p. 312), has
been included by Sava g e (2002, pp. 1095, 1159)
in the family Septalariidae Havlíček, 1960, while
Flabellulirostrum has been included in the family
Uncinulidae RzhonSnitSkaya, 1956.
The establishment of the Family Sapphicorhynchidae
SaRtenaeR, 2007 represented a second step in
the dismantling of the family Trigonirhynchiidae. The
genus Xahetomus SaRtenaeR, 2009 was added to the
type genus of the family.
There is no doubt that the breaking up of the
family Trigonorhynchiidae will be pursued, as the
introduction by GaRcía-Alcalde (2009) of the family
Iberirhynchiidae (+ subfamily Iberirhynchiinae), with
type genus Iberirhynchia DRot & WeStbRoek, 1966,
and the subfamily Myrmirhynxinae, with type genus
Myrmirhynx Havlíček, 1982, indicates. With the
exception of its inclusion in the family Oligorhynchiidae
CoopeR, 1956 by GaRcía-Alcalde (1998, p. 769),
Iberirhynchia has been initially assigned to the family
Trigonorhynchiidae, and maintained in it thereafter.
SaRtenaeR (2007, p. 43) criticized both this transfer
and the contents of this family. Both type genera were
included by Sava g e (1996, table 3, p. 256; 2002, p.
1056) in the subfamily Trigonirhynchiinae; Jin et
al. (1993, p. 54) had already assigned Myrmirhynx
to the same subfamily. All this explains why, in the
following comparison, the definition of the family
Trigonirhynchiidae will essentially rest on the definition
of its type genus, Trigonirhynchia fallaciosa.
The family Inaequalibellirostridae differs from the
family Trigonirhynchiidae in an outline that is constantly
subtrigonal (subtrigonal to subrounded in the family
Trigonirhynchiidae); a right angle triangle profile (half
an ellipse to half a circle in the globular to subcuboidal
family Trigonirhynchiidae); a convexo-concave ventral
valve; a deeply serrate commissure; a deeper sulcus;
higher fold and tongue; a lower number of higher and
sharper costae; the absence of parietal costae (but
external median costae are lower, sometimes markedly,
than the others) on the slopes of the high fold; the
maximum thickness at front margin and corresponding
to the top of tongue (in the family Trigonirhynchiidae
the fold curves more or less abruptly and flattens
towards the middle part of the anterior commissure
that is located around one third, exceptionally one half,
of front below the maximum shell thickness); dental
plates separated from the wall by narrow umbonal
cavities as already mentioned by SaRtenaeR (2007, p.
42) [Remark: as mentioned by SaRtenaeR (2007, p.
42) serial transverse sections made from two topotypes
of Trigonirhynchia fallaciosa do not show the wide
umbonal cavities figured by Schmidt (1965, fig. 1, p. 4),
and duplicated by Schmidt in Schmidt & McLaRen
(1965, figs 428, 1f-q, p. H559), and Sava g e (2002, figs
710, 1e-n, p. 1053)]; a deep septalium ; and a smaller
(narrower and less elongate) ventral muscle field.
Inaequalibellirostrum n. gen.
Derivatio nominis
Inaequalis, is, e (Latin, adjective) = inequal; libellus, i
(Latin, masculine) = level; rostrum, i (Latin, neuter) =
beak. The name draws attention to the inequal level of
the middle and external median costae of the species
assigned to the genus.
Type species: Hemithyris Pareti de VeRneuil, 1850a.
Species assigned to the genus
Besides the type species, Rhynchonella inaurita from

the “Spiriferensandstein” of the middle Rhine valley
described by the SandbeRgeR brothers (1856, pp. 336-
338, 444, 459, 470-472, 474, 477, 541, pl. XXXIII,
figs 5, 5a-c) is assigned to the genus. Three specimens
of the species are figured (Fig. 2). Some Lower
Emsian (“Untercoblenzschichten”) forms identified
as daleidensis in the regions E (Antweiler) and SE
(Stadtfeld, Oberstadtfeld) of the “Eifelkalkmulden”
probably belong to the genus. They are very close to
inaurita, but somewhat smaller. On account of lack of
material, the author is compelled to go no further in the
matter.
It has been mentioned above that the confusion of R.
inaurita with various species (Terebratula livonica as
understood at that time, T. Huotina, and T. Daleidensis)
was unacceptable as were the wide stratigraphic
range and geographic distribution that such confusion
implied.
The name inaurita chosen by the SandbeRgeR
brothers indicate that the absence of ears (read:
lunulae) in the dorsal valve (“Die Rückenklappe ist
gänzlich ohne Oehrchen”) is a major character of this
description. They also considered the lunulae observed
Fig. 2 –
Terebratula Daleidensis and related species
Inaequalibellirostrum inauritum (SandbeRgeR, G. & F.,1856). Dorsal, ventral, anterior, posterior, and lateral views
of three specimens. Topotypes (largo sensu) A-C, IRScNBa12714-12716. Speckgraben im Feisternachttal near Vallendar,
topographic map 1: 25.000 Bendorf, middle Rhine valley, Vallendar Substage (upper Lower Emsian).
65
on both valves of Rhynchonella Pareti as the main
difference between the two species, but expressed as
such, this is only partly true. As a matter of fact, ears
are present in R. inaurita, although not as strongly
marked as in R. Pareti. This is acknowledged by the
SandbeRgeR brothers, who stated that the dorsal
valve is “gänzlich ohne Oehrchen oder nur sehr kurz
geöhrt”.
GoSSelet (1887), the first author to accept that R.
inaurita could be a valid species (see above) reached such
a conclusion in examining SchnuR’s (1853, pl. XXII,
figs 1a-c) figures of R. Daleidensis that he considered
different from the original ones of RoemeR (1844, pl.
I, figs 7a-c). It is easy to find in the literature specimens
identified as (Terebratula, Rhynchonella) daleidensis
that belong to R. inaurita, e.g. QuenStedt (1871, pl.
42, fig. 59; “Grauwacke” Daun, Eifel area) duplicated
by RoemeR (1876, pl. 23, fig. 7), GüRich (1909, pl.
45, figs 6a-d = pl. 33, figs 5, 5a-c in SandbeRgeR, G.
& F., 1856).
Figures of three specimens [topotypes (largo
sensu) IRScNBa12714-12716) from the Vallendar
Substage (upper Lower Emsian) at Speckgraben im

66 Paul SARTENAER
“Feisternachttal” near Vallendar (Middle Rhine valley)]
are shown on Fig. 2; they are part of a collection of
more than one hundred specimens made by Dr. K.-W.
Wenndorf, and were generously presented to the
author. Inaurita is restricted to the Vallendar Substage.
Some isolated embedded valves suggest that it could
also be present in the lower Upper Emsian Emsquarzit,
Lahnstein Substage, and in “Hohenrheiner Schichten”
of the middle Rhine valley, but there is no conclusive
evidence for it.
Only two German geologists identified as R.
inaurita specimens collected from two very different
stratigraphic levels in the Middle Rhine valley. These
are MauReR (1886, pp. 27, 37, 53), and FuchS (1899, p.
68), who reported R. inaurita from the “Cultrijugatus-
Stufe” (= Stufe VIII = uppermost Emsian) at
Laubach and Michelbach), and the basal Hercyniae
Zone (middle Lower Emsian) at Bellsgraben in the
Lorelei area, respectively. MauReR (1896, p. 658)
even suggested that R. inaurita could eventually be
considered as the dominant species (“vorherrschende
Art” of what he defines as the lower subdivision [=
Stufen I (“Taunusquarzit”), II (“Hunsrückschiefer”),
and III (“untere Grauwacke”)] of the Lower Devonian
on the east of the Rhine, and R. daleidensis as the
dominant species of the upper subdivision [= Stufen
IV (“Haliseritenschiefer”), V (“Coblenzquarzit”), VI
(“Chondritenschiefer”), VII (“Hohenrheiner-Stufe”),
and VIII “Cultrijugatus-Stufe”)]. The pertinence
of these identifications cannot be assessed by the
author.
The “very abundant” Devonian form from
Devonshire identified as Rhynchonella inaurita by
DavidSon (1870, pp. 72, 73-75, 78, 79, 80, pl. V, figs
1-3; 1881, pp. 336, 340, 341, 343, 351-353, pl. 38, fig.
21?, 35, 35a,b), and the Frasnian (sic!) form from NW
Poland (borings) identified as Camarotoechia (C.) ex
gr. inaurita by ŁobanowSki (1968, pp. 768, 774, 775,
783, pl. II, figs 4-5) do not belong to that species.
Description
Shell medium- to medium-large-size, profile a rightangle
triangle, strongly dorsibiconvex, and width
always the largest dimension. Shell outline subtrigonal.
Maximum shell width located anteriorly. Ventral
lunulae much lower than the dorsal ones on account
of the convexo-concave ventral valve; these posterolateral
concavities, also called ears, may extend as far
as mid-length and are separated from the flanks proper
by ridges that are well pronounced in the umbonal
region. One, exceptionally two, faint costae present in
the lunulae.
Anterior and antero-lateral commissures deeply
serrate, postero-lateral commissures only slightly.
Apical angle moderately wide. Sulcus and fold strongly
marked, start from beaks. Ventral flanks narrow. Beak
pointed, projecting, slightly to strongly incurved.
Interarea very narrow. Deltidial plates short. Sulcus
deep, widening rapidly, wide at front, with flat bottom,
extended dorsally as a high and clearly delineated
tongue with variable outline (trapeze, semicircular
arch, and gothic arch). Tongue high, recurving slightly
to strongly posteriorly in its uppermost part in the
largest specimens. Crest of tongue variously (slightly
to strongly) curved.
Dorsal valve very high, thickest at front margin,
vertical or almost vertical in the umbonal area.
Fold very high, rapidly gaining in height anteriorly,
sometimes a sharp rise near front. Slope of flanks
sharply interrupted postero-laterally by the lunulae,
abrupt antero-laterally. Costae few, simple, regular,
high, wide, angular (acute, sharp), starting from beaks.
No parietal costae. Shell thin. Teeth stout, wide, short,
cyrtomatodont, entering the dental sockets vertically in
serial transverse sections. Denticula well developed.
Dental plates slender, subparallel, separated from the
wall by wide umbonal cavities. Delthyrial cavity wide.
Hinge plate undivided. Outer hinge plates narrow.
Septum extending until mid-length. Septalium wide,
deep, with variable shape, median ridge sometimes
present on bottom of septalium. Connectivum covering
anterior part of septalium. Crura long, raduliform,
aliform in section in their proximal part, walkingstick-shaped
in their distal part, where they are curved.
Dental sockets shallow, bottom wrinkled, inner socket
ridges high. Muscle fields slightly impressed.
Comparisons
Inaequalibellirostrum can easily be separated from
Oligoptycherhynchus by a right-angle triangle profile,
a subtrigonal outline, well marked lunulae, a slightly
wider apical angle, a slightly deeper sulcus, a tongue
with variable outline, a higher fold with (sometimes) a
sharp rise near front, acute and slightly higher costae,
a different number of median costae, the absence of
parietal costae; a deeper septalium; longer crura; and
teeth entering the dental sockets vertically in serial
transverse sections (laterally in Oligoptycherhynchus).
Representatives of the genus Inaequalibellirostrum
may also reach a larger size.
Inaequalibellirostrum differs from Sapphicorhynchus
SaRtenaeR, 2007 by many features, the major
ones being a right-angle triangle profile, a subtrigonal
outline, well marked lunulae, a smaller apical angle, a

deeper sulcus, a higher tongue with variable outline,
a higher dorsal valve thickest at front, a higher fold
with (sometimes) a sharp rise near front, acute costae,
4 5 7
a different general costal formula [ ; to for I.
3 6 8
5 − – 6 0 – − 1 1 − – 1 6 − 8
pareti against ; 0 to , and ;
6 – 8
for
46
−–
5 0 – −1
1 1 −–
1
7 −–
9
S. sappho].
Inaequalibellirostrum pareti (de VeRneuil, 1850a)
Fig. 3
The following generic assignments were given in
succession to the species: Terebratula MülleR,
1776, Rhynchonella FiScheR de Waldheim, 1809,
Camarotoechia Hall & ClaRke, 1893, “Camarotoechia”,
Trigonirhynchia CoopeR, 1942, Stegerhynchus
FoeRSte, 1909, and Oligoptycherhynchus
SaRtenaeR, 1970.
In more recent time, not to mention the assignment
to Trigonirhynchia CoopeR, 1942 by WeStbRoek
(1967, fig. 7, p. 8, pp. 9-10, 20-21, fig.19, p. 21, fig.
21, p. 22, figs 26-28, p. 26, p. 27, fig. 27, p. 27, pp.
28-29, p. 30, fig. 32, p. 30, fig. 35, p. 32, pp. 34, 66,
69, 70, pl. III, figs 1, 7, 7a, pl. V, figs 1, 2, 2a, 3, pl. VI,
figs 1a-c, pl. VIII, figs 1a,b, 2, 3) with a considerable
stratigraphic range (Upper Siegenian - Lower Eifelian),
and questionably to Stegerhynchus FoeRSte, 1909
by GaRcía-Alcalde et al. (1979, fig. 16, p. 27), the
species has since 1984 continuously been mentioned as
Oligoptycherhynchus pareti.
Lectotype
The only specimen figured by de VeRneuil (1850a,
pl. III, figs 11a,b) is here designated as the lectotype
of the species. de VeRneuil does not indicate from
which of the two localities specifically mentioned in
his description of the species (Sabero in the Province
of Leon, and Ferroñes in the Province of Asturias) this
specimen comes from.
The author is inclined to believe that Colle, referred
to by de VeRneuil, 1850a, p. 164 as “the richest locality
of the Sabero district”, is the locality from where the
specimen has been collected. The species is particularly
abundant there, and de VeRneuil (1866, p. 11) wrote
that he found it «pour la première fois dans les calcaires
dévoniens inférieurs de Sabero».
Description and Remarks
Besides the original description of the external characters
by de VeRneuil (1850a, p. 177, pl. III, figs 1a,b) a
Terebratula Daleidensis and related species
67
more detailed one has been given by OehleRt (1884,
pp. 415-416, pl. XIX, figs 2, 2a-i). The description by
Schumann (1965, p. 93, fig. 2, p. 95, p. 96, pl. 4, figs
10a-c as Camarotoechia daleidensis) includes internal
characters.
BaRRoiS’s (1889, pp. 84-85, pl. V, figs 3a-c)
description of the alleged Armorican Rhynchonella
Pareti is almost a word for word transcription of
OehleRt’s description of the Spanish species.
No comparison to R. subpareti OehleRt, 1884 is
attempted; R. subpareti is only listed (p. 328) in “la
faune coblenzienne des calcaires de Bretagne”.
The description that follows refers only to specific
characters in need of further elaboration. Length and
thickness similar, but thickness is usually slightly
larger (
t
between 0.81 and 1.23, mostly between 0.85
l
and 1.10; l between 0.74 and 0.87;
t
w
w
between 0.66
and 0.94). Width of sulcus at front between 63 and
85%, mostly between 70 and 80%, of shell width.
Maximum shell width between 60 and 71% of shell
length anterior to the ventral beak. Maximum thickness
of dorsal valve, and thus of shell, between 81 and 70%
(exceptionally less) of shell length anterior to ventral
4
beak. median costae (very exceptionally
5
or even
3
4
6
4
7
8
), to (exceptionally ) lateral costae; mere
5
5
8
9
undulations of the postero-lateral commissures are not
included in the counting. Median costae 3 to 4 mm wide
near front. External costae of fold, distinctly lower than
the others, and the ventral internal lateral costae, higher
than the others, could rigorously be called parietal, but
slopes of the fold are free of costae, as suggested by the
name of the species (paries, etis, Latin = wall, flank).
Apical angle between 103 and 115°, mostly around
105°. Transverse serial sections from one topotype
(IRScNBa12713) are shown in Fig. 3. Sections made
in five more specimens indicate that the septalium is
either cupula-, amphora- or tumbler-shaped.
Excellent transverse serial sections have been made
by DRot (1964, fig. 75, p. 181) from a specimen from
the Province of Asturias (no precise locality given), and
by Schumann (1965, fig. 22, p. 95 as Camarotoechia
daleidensis) from a specimen from the type area
(Aguasalio syncline). Another transverse section and
excellent reconstructions of the internal characters of
the species have been figured by WeStbRoek (1967,
fig. 7, p. 8, fig. 32, p. 30, fig. 35, p. 32).
Collections mentioned by Alva R e z (1990, pp. 306-
307) at two localities of the Province of Leon (Colle E
of Boñar, and Villayandre S of Crémenes) were loaned

68 Paul SARTENAER

to the author by Prof Fernando Alvarez; they allowed
assessing the variability, the ontogenetic development,
and the number of costae of the species.
Comparisons
Various characters make pareti distinct from daleidensis:
a right-angle triangle profile, a subtrigonal outline, well
marked lunulae, a smaller apical angle, a lower ventral
umbonal region, a projecting ventral beak, flat ventral
flanks, a slightly deeper, better marked, and often wider
sulcus, a tongue with variable outline, a relatively higher
dorsal valve, always thickest at front, a higher fold with
(sometimes) a sharp rise near front, a different general
4
costal formula [
4
;
5
to
7
(exceptionally
8
) for
3 6
8
9
4
7 – − 9
pareti against ; 0; for daleidensis], external
3
6 – −10
10
costae of fold distinctly lower than the others.
Inaequalibellirostrum inauritum can easily be
separated from I. pareti by a larger size, a deeper
sulcus, a higher fold, a higher tongue, and a slightly
7 9
higher number ( to
9
) of lateral costae. Although
8
10 10
variable in both species, the tongue outline of I.
inauritum is more often a gothic arch than in I. pareti.
In extremely rare specimens, the external costae of fold
of I. inauritum are not only lower than the others as in
I. pareti, but they are significantly lower, and have to
be called parietal.
Stratigraphic range and geographic distribution
de VeRneuil (1866, p. 11), who previously (1850a,
1850b) had only indicated a Devonian age for the
species, was more precise when he declared that
he found the species he established in the “calcaires
dévoniens inférieurs de Sabero”.
In the type area (Province of Leon),
Inaequalibellirostrum pareti is found in the part of the
La Vid Group corresponding to the upper part of the
Valporquero Formation and the Coladilla Formation,
i.e. in the middle Emsian Faunal Intervals 11 to 13
introduced by GaRcía-Alcalde (1994). It means, in
terms of the conodont zonation, the middle and upper
parts of the undifferentiated Polygnathus laticostatus/
inversus Zone + the lower half of the P. serotinus Zone
[GaRcía-Alcalde in GaRcía-Alcalde et al., 1979,
fig. 16, p. 27; in TRuyolS et al., 1990, fig. 1, p. 14;
1994, fig. 2, p. 78; 1995, fig. 6, p. 21; 1996, fig. 2, p.
Fig. 3 –
Terebratula Daleidensis and related species
Inaequalibellirostrum pareti (de veRneuil, 1850). Camera lucida drawings of serial transverse sections; figures are
in mm forward of the ventral umbo. Topotype IRScNBa12713. Measurements: length = 17.9 mm; width = 19.2 mm;
thickness = 17 mm. (opposite page)
69
60; 2001, fig. 1, p. 548), and Alva R e z & BRime (1990,
table 1, p. 15)].
Remark: Spanish geologists prefer to include Faunal
Intervals 11 to 13 in the lower part of the Upper Emsian,
because they subdivide the Emsian into two parts: a
Lower Emsian (= most of Pedrosa Formation), and an
Upper Emsian (= uppermost part of Pedrosa Formation
+ Valporquero Formation + Coladilla Formation + most
of the Santa Lucia Formation).
Is pareti present outside the type area?
According to the literature, the species is present in
three countries: France (Armorican Massif, Ardennes,
Belfort Territory in the “Département de la Franche-
Comté”, Pyrenees), Spain (Cantabrian Cordillera,
Central Iberian Zone), and Turkey (Asia Minor), and
ranges from the Gedinnian to the Lower Eifelian. This
has little interest.
The species has been mentioned in the Asturian
Devonian by various authors since de VeRneuil (1850a,
pp. 160, 177; 1850b, table, p. 780, p. 784; 1866, p. 11),
and de VeRneuil & BaRande (1855, p. 1016).
BaRRoiS (1882, pp. 267-268, 473-475, table, p. 503,
table, p. 518) collected the species from the “calcaire(s)
de Nieva” (or “schistes et calcaires de Nieva” or “zône
de Nieva”) at various localities of the Asturian coast,
and attributed to them an early Coblencian age (i.e. late
Siegenian and early Emsian). The Nieva Formation is
considered nowadays of late Lochkovian- early Pragian
age, thus indicating a considerably older age than the
beds containing pareti in the type area.
From the beginning the species has been mentioned
in Puerto del Ciervo and Chillón in the Sierra Morena
(Central Iberian Zone) by de VeRneuil in de VeRneuil
& BaRRande (1855, p. 1016; 1866, p.11). Recent
investigations by PaRdo & GaRcía-Alcalde (1984a,
p. 83; 1984b, p. 475; 1994, fig. 2, p. 155; 1996, fig.
3, p. 75, p. 79) and PaRdo-AlonSo (in GaRcía-
Alcalde et al., 2000, fig. 4, p. 138) have indicated
that Oligoptycherhynchus cf. pareti, O. gr. pareti and
O. aff. pareti are part of a middle Emsian fauna in the
Central Iberian Zone and in the northern part of the
Ossa-Morena Zone.
The occurrence of pareti in the Ardennes rests on
BaRRoiS’s (1882, p. 512) statement that the species was
common to the “Nieva limestone” and the “Montigny
fauna”. BaRRoiS, however, had in mind daleidensis
that he considered, as previously indicated, as closely

70 Paul SARTENAER
related to pareti, livonica, and inaurita. The presence of
daleidensis in the late Siegenian Montigny Greywacke
had already been mentioned by French and Belgian
geologists, especially by GoSSelet in many of his
papers published between 1863 and 1880.
RouSSel (1904, p. 18), and ASSelbeRghS (1926,
p. 72) mentioned the presence of the species in the
Pyrenees and in the Belfort Territory (Parisot-Chevillard
collection) respectively.
The presence of pareti in the regions just mentioned
must be disregarded, but what about the Armorican
Massif, where its presence has been widely accepted
during the second half of the 19 th century?
In his original description of Hemithyris Pareti,
de VeRneuil (1850a, pp. 159-162, 177, pl. III, figs
1a,b) made a distinction between this Devonian species
“propre à l’Espagne” (Asturias and Leon) and a
species “presque identique” from Viré-en-Champagne
(“Département de la Sarthe”, Laval Syncline). The
same year (1850b, table, pp. 780-781, pp. 784-785,
786 as Terebratula Pareti), however, he abandoned
this distinction, and declared the species present
not only in the Cantabrian Cordillera, but also in the
Sierra Morena, at Viré-en-Champagne, Brûlon, and
Joué (“Département de la Sarthe”, Laval Syncline),
and Néhou (“Département de la Manche”, Cotentin
peninsula, Lower Normandy); he even explicitely
mentioned the “Terebratula Pareti de notre pays”.
Pareti from the Laval Syncline, to which BaRRoiS
(1889, p. 85) gave the same qualification («type») as
to the Spanish species. OehleRt (1877, pp. 592, 601)
named Terebratula Pareti specimens collected from
the “Calcaire à Spirigera undata de la partie moyenne
du dévonien inférieur” (probably middle Pragian)
at la Baconnière, Saint-Germain, and Saint-Jean
(“Département de la Mayenne”); in 1884 (p.414) he
replaced this identification with T. cypris.
No pareti has been spotted in the various collections
from Néhou, where its presence has been indicated
by de VeRneuil (1850b), by OehleRt (1884, p. 416,
and since then by various authors. Outside of the
Laval Syncline and Néhou, pareti has been mentioned
thereafter from the “Rade de Brest”, and Erbray.
Pareti was recorded from the “Rade de Brest”
(Châteaulin Syncline, “Département du Finistère”)
by de TRomelin & LebeSconte [1876, p. 611 as
Rhynchonella Paretoi (R. Cypris, d’Orb.); Lower
Devonian], BaRRoiS¨ [1886, p. 691; 1899, p. 239,
«grauwacke du Faou» now considered as middle
Pragian-early Emsian in age, and «grauwacke du Fret»
(late Emsian in age). No available collection allows
substantiating these claims. It is doubtful that these
references could indicate the small “Rhynchonella”
cf. subpareti described by BRice (1980, pp. 238-239,
figs 58A, B, p. 237, fig. 59, p. 241, pl. 33, figs 6a, b;
1981, p. 197), and revived by MoRzadec et al. (1988,
fig. 7, p. 12-13; 1991, fig. 4, p. 907) from the Lower
Pragian of the “Pointe de l’Armorique” located in the
same syncline (see below); however, this age differs
from those just mentioned. Although BRice (1980,
pp. 238-239, figs 58A,B, p. 237, pl. 33, figs 6a,b),
who figured one specimen and made transverse serial
sections from two others, considered her identification
as questionable, it cannot be disregarded.
BaRRoiS (1887), after having mentioned the
presence of pareti in the “Faune d’Erbray” (p. 160
as Rhynchonella pareti), he (1889, pp. 84-85, 87,
table, p. 251, pp. 258, 261-262, 273, 328, pl. V, figs
3a-c), described R. Pareti from the Erbray Limestone
at Erbray [Saint-Julien-de-Vouvantes Syncline,
«Département de Loire-Inférieure” now “Département
de la Loire-Atlantique”)]. The species had been already
mentioned there by Cailliaud (1861, p. 333 as
Terebratula Pareti or hemithyris) and de TRomelin &
LebeSconte (1876, pp. 606-607) [Remark: the word
“unknown” used by BaRRoiS (1889, p. 261) to qualify
Cailliaud’s Terebratula Pareti from the Erbray
Limestone is misleading. Does it mean that he did not
find the specimen(s) in Cailliaud’s collection that he
had at his disposal? Otherwise, it is a contradiction,
because he described Rhynchonella Pareti from the
same limestone].
BaRRoiS’s description indicates, on one hand, that
he had a sizeable collection at his disposal, and on the
other, that more than one species was present in the
material on hand, among others the “petites coquilles
trigones” that he considered as juveniles.
The only figured specimen is easily separable from
pareti by its proportions, in particular by its smaller
thickness. It lacks the following characteristic features
of pareti, i.e. the right-angle triangle profile, the wellmarked
lunulae, the very high fold, the usual number
4
4
( ) of median costae (although is mentioned by
3
3
BaRRoiS as the general ratio), the external costae of
fold markedly lower than the others, and the high
median costae. The present author is inclined to believe
that this specimen belongs to a species of its own. This
seems to be implied by BaRRoiS himself, who notes
(1889, p. 85) that Rhynchonella Pareti from the Erbray
Limestone, that he incorrectly dates as Gedinnian (late
Pragian – early Emsian is the prevailing age nowadays),
“se distingue un peu, de mes types de l’espèce, de la
Sarthe [subpareti, according to the pesent author] et

de l’Espagne [pareti], par sa forme plus transverse,
moins haute à l’état adulte”. This opinion is supported
by a small collection of the “Musée des Sciences “
in Laval, comprising four specimens from the Lower
Devonian of la Jaillerie near la Baconnière in the
nearby “Département de la Mayenne”, all of them of
the same size as the specimen figured by BaRRoiS, and
4
with median costae.
3
After 1889 mentions of pareti disappeared almost
completely from the French literature related to the
Armorican Massif, with the exception of indication
of its presence in a given area (Couffon, 1925, p.
45 as Rhynchonella pareti, “Département de Maineet-Loire”)
and the confirmation of its occurrence
in various localities (Péneau, 1929, p. 263 as R.
pareti, Erbray, Lower Devonian; Comte, 1938, p. 59,
Brittany, Coblencian). In short, pareti is not present in
the Armorican Massif.
What about subpareti, which is a poorly known
Middle Pragian species from the Armorican Massif? Its
name alone imposes a comparison with pareti.
OehleRt (1884) gave a new and more complete
description of the Spanish pareti (pp. 415-416, pl. XIX,
figs 2a-i as Rhynchonella Pareti), the presence of which
he still accepts in Néhou (p. 416), and established (pp.
412, 414-417, pl. XIX, figs 3, 3a-e) Rhynchonella
subpareti OehleRt, 1884 that he had once considered
«comme une simple variété de R. Pareti», and that
he now sees as a «modification locale de la R. Pareti
d’Espagne, mais dont les caractères sont suffisamment
fixés pour qu’il soit nécessaire de lui donner un nom
distinct» or a «forme représentative de la R. Pareti
d’Espagne, mais qui, pour ses caractères constants,
mérite d’être désignée par un nom distinct» or as
offering «des modifications suffisamment constantes et
distinctes du type d’Espagne, pour qu’il soit nécessaire
de la mettre à part». This phraseology presupposes the
extreme and not acceptable plasticity of a «type» similar
to the one of R. livonica advocated by Kay S e R (1871b)
(see above): « ces espèces [R. cypris, Pareti, subpareti,
livonica, nympha, pseudolivonica, etc.] ne sont sans
doute qu’un même type, modifié dans le temps ou dans
l’espace». A similar plasticity is advocated by BaRRoiS
et al. in GoSSelet et al. (1922, p. 97) for a group
including R. nympha, R. daleidensis, R. livonica, R.
sub-livonica, R. Pareti, and R. sub-pareti (see above).
The two specimens of Rhynchonella subpareti
figured by OehleRt (1884, pl. XIX, figs 3, 3a-e; the
specimen of figs 3, 3a,b is here formally designated
as the lectotype of the species) went astray fide BRice
(1980, p. 238). According to DRot (1964, p. 177) and
Terebratula Daleidensis and related species
71
DRot & L’HotellieR (1976, p. 268), OehleRt’s
figures of Rhynchonella subpareti represent a juvenile
form (figs 3c-e) with the aspect of R. cypris d’ORbigny,
1847, which could be “Camarotochia paretiformis”
DRot, 1964 from the Lower Gedinnian of the Dra
Plains (Anti-Atlas, Morocco). Very few specimens of
this taxon collected at that time are available, although
OehleRt (1884, p. 415) stated that the species he
established was quite frequent in the Devonian outcrops
of the “Département de la Sarthe”. BRice in MoRzadec
et al. (1988, p. 52) also reminds that OehleRt collected
the species from the very fossiliferous Pragian (Saint-
Céneré Formation) shales and limestones of the old
quarry La Roussière SW of Saint-Germain-le-Fouilloux
(“Départemaent de la Sarthe”).
The four localities mentioned by OehleRt in 1884
(Saint-Céneré in the “Département de la Mayenne”,
and Viré, les Courtoisières, and Vieux-Michel in the
adjoining “Département de la Sarthe”, all localities in
the Laval Syncline) were dropped by OehleRt (1886,
p. 23). Instead a fifth locality of the same syncline,
Sablé (“Département de la Sarthe”), was added.
BaRRoiS (1889, p. 328) did not discuss Rhynchonella
subpareti that he only listed in the “faune coblenzienne
du calcaire de Bretagne”.
As a matter of fact, subpareti never gained wide
acceptance in the literature related to the Armorican
Massif. Except for the mentions by Reed (1922, p. 96)
and BaRRoiS et al. in GoSSelet et al. (1922, p. 97)
(see above), and by Giovannoni & ZanfRa’ (1979, p.
214 as Camarotoechia subpareti), subpareti has been
reported by Couffon [1925, p. 78 as Rhynchonella
subpareti, “Département de Maine-et-Loire”, Lower
Givetian (sic!)], Péneau [1929, pp. 84, 231, 263
as R. sub.Pareti, Angers-Saint-Julien-de-Vouvantes
Syncline, “Département de Maine-et-Loire”, Calcaire
de Vern (Middle Siegenian, now Upper Lochkovian-
Lower Pragian)], and Pillet (1953, p. 17, Calcaires de
Vern-d’Anjou).
Still, as previously indicated, subpareti popped up
again in another part of the Armorican Massif when
BRice (1980, 1981) described “Rhynchonella” cf.
subpareti from the Lower Pragian of the “Pointe de
l’Armorique” (Châteaulin Syncline, “Rade de Brest”,
“Département du Finistère”). The stratigraphic range
of this species was again given by MoRzadec et al.
(1988, 1991). BRice (1980, p. 239) stated that “R.” cf.
subpareti seemed to belong to the same genus as “R.”
nympha that had still to be defined. Consequently BRice
(1981, pp. 195, 197, p. 214, figs 1A, B, p. 196, fig. 7,
p. 213, pl. 25, figs 1a-c, 2a-d, 3a,b, 4, 5) proposed the
genus Stenorhynchia BRICE, 1981 with type species

72 Paul SARTENAER
Terebratula nympha BaRRande, 1847 and included in
it the varieties proposed by BaRRande. At the same
time, she assigned OehleRt’s Rhynchonella subpareti
to it, and described as nympha an Upper Emsian small
form from the la Lézais trench in the Ménez-Bélair
Syncline (“Département d’Ille-et-Vilaine”) assigned
by SaRtenaeR (2009, p.34) to the genus Xahetomus
SaRtenaeR, 2009. MoRzadec et al. (1981, fig. 4, p. 12,
fig. 5, p. 14, pp. 16-17) and MoRzadec (1983, p. 276,
fig. 6, p. 286) confirmed the presence of nympha in the
Upper Emsian of the la Lézais trench, and mentioned it
also from beds of the same age in the Reun ar C’Hrank
en Lanvéoc section (“Rade de Brest”), which LaRdeux
& MoRzadec (1979, pp. 14, 17) had already reported
Stenorhynchia nympha from.
This brings up questions regarding the presence of
the Bohemian species in the Armorican Massif and the
contents of the genus Stenorhynchia. The presence of
nympha in the Armorican Massif, covering the form
identified and figured as such by BaRRoiS (1889) and
also various other forms, has been accepted by many
authors, e.g. Cailliaud (1861, p. 332), de TRomelin
& LebeSconte (1876, pp. 606-607), Kay S e R (1878,
p. 143: Pareti “gehört wahrscheinlich zu nympha”),
OehleRt (1884, p. 416; his opinion has been mentioned
above), FRech (1887, p. 410), BaRRoiS [1889, pp. 86-
87, table, p. 241, p. 328, pl. V, figs 2a-e; nympha, of
which he had only six specimens, is figured for the first
time and considered distinct from Pareti present in the
same limestone (Erbray Limestone)], MauReR (1896,
p. 659), Venyukov (1899, p. 157), Scupin (1906, pp.
236-238, table, p. 302), Reed (1922, p. 96; refers to
nympha “as figured by BaRRoiS”), Péneau (1929, p.
230, table, p. 263), Le MaîtRe (1934, pp. 210-211,
table, p. 229, pp. 230, 235, 239; 1944, pp. 11, 12, table,
p. 20, pp. 47, 48), Comte (1959, p. 276), Havlíček
(1961, p. 87), Pillet (1962, p. 49), DRot (1964, p.
105). Maillieux (1931, pp. 21, 24) also indicated
the presence of nympha in the Armorican Massif and
envisaged the possibility that it belongs to Pareti, and,
as a consequence, falls into synonymy of daleidensis),
L’HotellieR-Videau, who devoted her PhD
(1970) to the rhynchonellids from some outcrops in the
southeastern part of the Armorican Massif, discussed
neither R. pareti nor R. subpareti, but described eight
distorted specimens from existing collections from the
Erbray Limestone as Stegerhynchus? nympha and S.?
cf. nympha. She figured three specimens representing
two, probably three, taxa, none of them approaching R.
Pareti or R. nympha figured by BaRRoiS (1889, pl. V,
figs 2a-e, 3a-c).
The Upper Emsian S. nympha from la Lézais is also
not identical with the Pragian S. nympha from Bohemia
(age fide Havlíček, 1961, pp. 85, 87) [Remark: the
species described as S. nympha by BRice (1981) is
completely different from Rhynchonella nympha
described by BaRRoiS (1889)].
Various characters make the Bohemian nympha
distinct from the Armorican nympha: a ventral valve
almost completely excavated by the sulcus, a wide and
shallow sulcus with flat bottom, a larger number of
lateral costae (8 to 13), the almost systematic presence
of parietal costae, and the absence of a connectivum
[Havlíček (1961, p. 86) writes: “Das Septalium
wird von ventraler Seite durch zwei dünne Fortsätze
eingeengt, die den oberen Septaliumränder aufsitzen
und deren freien Enden gegeneinander orientiert
sind. Diese Fortsätze berühren sich jedoch niemals”;
(see also fig. 29, p. 86 of Stegerhynchus nympha)].
[Remark: GaRcía-Alcalde in TRuyolS-MaSSoni &
GaRcía-Alcalde (1994, p. 232), taking into account
a personal communication from BRice, declares
that transverse serial sections made by her from two
Bohemian specimens presented by Havlíček have
shown a delicate tectiform connectivum in the anterior
part of the septalium].
Nympha from la Lézais does not belong either to
the genus Stenorhynchia. “Rhynchonella” cf. subpareti
from the “Pointe de l’Armorique” also cannot be
assigned to the genus Stenorhynchia, and the assignment
of Terebratula Nympha pseudo-livonica BaRRande to
Stenorhynchia is also disputable.
GaRcía-Alcalde in TRuyólS-MaSSoni &
GaRcía-Alcalde (1994, pp. 232-233) included
BRice’s Upper Emsian species from la Lézais in
Stenorhynchia briceae GaRcía-AlcaldE, 1994, a
frequent species from the upper third of the La Ladrona
Formation and the basal part of the Aguión Formation of
the Cantabrian Cordillera, i.e. from the middle Emsian
Faunal Intervals 9 to basal 12 of GaRcía-Alcalde
(Remark: these Faunal Intervals are put in the Upper
Emsian by Spanish geologists because they subdivide
the Emsian into a Lower Emsian restricted to the lower
half of the La Ladrona Formation, and an Upper Emsian
corresponding to the upper half of that formation + the
Aguión Formation + most of the Moniello Formation).
This substitution, accepted by BRice (2000, p. 15), does
not make the generic assignment more acceptable.
Terebratula nympha, a characteristic species of the
Konĕprusy (Pragian) Limestone, has successively been
assigned to the following genera (in parentheses are
indicated the main periods): Rhynchonella fiScheR
de Waldheim, 1809 (1856-1963), Camarotoechia
Hall & ClaRke, 1893 (1934-1990), Nymphorhynchia

RzhonSnitSkaya, 1956 (1956-1987), Stegerhynchus
FoeRSte, 1909 (1961-1969), Stenorhynchia BRice,
1981 (1979-2000); assignments to Ancillotoechia
Havlíček, 1959 (1975), Felinotoechia Havlíček, 1961
(1982), and Microsphaeridiorhynchus SaRtenaeR,
1970 (1983) remain exceptions. Nympha is not present
in most of the regions of the world where it has been
mentioned; it is one of the various Lower Devonian
Bohemian species that have been unduly “exported”.
The genus Stenorhynchia has followed the
unfortunate usual pattern, shared with many genera,
in being as from its establishment overloaded with
species that are foreign to it. Today twenty-two species,
subspecies, and forms in open nomenclature have been
assigned to the genus. For example, neither the Upper
Emsian S. briceae already mentioned nor the five
small middle and late Silurian Bohemian species and
subspecies of Stenorhynchia described by Havlíček
in Havlíček & ŠtoRch (1990, pp. 28, 30, 38, 40, 42,
142-144, pl. XXXVII, figs 6a-c, pl. XLII, figs 1a-c,
2a-c, 4a-c, 5a,b, pl. XLIII, figs 4a-c, 5a,b, 6a-c, 7a-d,
8a-c, 9) belong to it.
A middle Silurian to early Zlichovian age advocated
for Stenorhynchia by Havlíček (1992, table 1, p.
56, p. 80) due to the extended (in relation with the
previous Pragian range) stratigraphic range he gives
to S. nympha, and to his questionable assignment,
in 1990 (in Havlíček & StoRch), of new Silurian
species to the genus [Remark: Chlupáč et al., 1972,
pp. 122, 148, 171 as Nymphorhynchia aff. nympha, and
Chlupáč et al., 1979, p. 146 as N. cf. nympha, already
indicated that Stenorhynchia nympha could be present
in the Lower Lochkovian and in the Upper Zlichovian
respectively].
Other stratigraphic ranges are also found in the
literature, e.g. Ludlow-Přídolí in the European Province
(Rong et al., 1995, appendix 2, p. 60) (it means that the
stratum typicum of the type species is excluded), and
those including the Emsian species incorrectly assigned
to the genus: Pragian-Emsian (Sava g e, 1996, table 3, p.
256), uppermost Lochkovian-Pragian-Emsian (BRice
in BRice et al., 2000, fig. 1, p. 68), and Upper Silurian
(Ludlow) to Lower Devonian (Emsian) (Sava g e, 2002,
p. 1062).
Compilation of the literature suggests long range
of the genus Stenorhynchia (Middle Silurian - Upper
Frasnian) and wide geographic distribution, since it has
been reported from Algeria (various regions), Armenia,
Belgium, Bohemia, Canada (various regions), Carnic
Alps, England (various regions), France (various
regions), Germany (various regions), Kazakhstan,
Lettonia, Libya (Fezzan), Mauritania (Zemmour),
Terebratula Daleidensis and related species
73
Mongolia, Morocco (various regions), North America
(various States), Podolia, Russia (many regions),
Spain (various regions), Tadjikistan, Turkestan, Turkey
(Bithynia), Uzbekistan. It is clear that the contents of
the genus are unduly inflated and that, in particular, the
world distribution of its type species is not acceptable.
According to the author, the genus Stenorhynchia is
restricted to the Pragian, with the suspicion that further
investigations will allow to restrict its stratigraphic
range even more within that stage.
Subpareti was “imported” into the Iberian
Cordillera by Comte (1959, p. 282, Coblencian, as
Camarotoechia subpareti), in the Ossa-Morena Zone
by PaRdo AlonSo & GaRcía-Alcalde (1996, p. 79
as Oligoptycherhynchus gr. subpareti, Upper Emsian),
and, as from 1992, as Stenorhynchia subpareti in the
Upper Pragian and Emsian Intervals 7 to 10 of the
Cantabrian Cordillera by GaRcía-Alcade (1992,
fig. 4, p. 59; 1994, fig. 2, p. 78; in TRuyólS-MaSSoni
& GaRcía-Alcalde, 1994, fig. 2, p. 223, pp. 232,
233; 1996, fig. 2, p. 60), and by GaRcía-Alcalde &
TRuyólS-MaSSoni (1994, p. 86, fig. 2, p. 87).
In the absence of any description of the Spanish
subpareti, it is not possible to accept the presence of
the Armorican species in any region of Spain. GaRcía-
Alcalde (1997, p. 243; 1998, p. 243) found S.
subpareti in Lower Emsian beds from the Dra Plains
(Anti-Atlas, Morocco); it is only mentioned here for
the sake of completeness.
How does subpareti compare with the Spanish
pareti? Subpareti did not benefit of a clear-cut
definition when it was established. As already
indicated, its original description is obscured by sybillic
phraseology. OehleRt (1884, pp. 414, 417) opposed
diminished (“amoindries”), attenuated (“atténuées”),
and exaggerated (“exagérées) specimens (or forms) of
subpareti and pareti to “typical” specimens (or forms)
or “normal types” of both species. Are we faced with
the ontogenetic development of one species or with
the transition from one species to another, an opinion
evoked by OehleRt himself (p. 414)? In any case the
wording used for establishing subpareti is ambiguous.
In his original description of subpareti, which is
more a comparison with pareti than the description of
an independent species, OehleRt (pp. 416-417) states
that the new species is smaller, less cuneiform, has a
more clearly marked outline, a more globular profile,
weaker and less angular costae, and less pronounced
lunulae. He also stresses that the relative depth of
valves and the development of the lateral and median
parts are different. The occasional secondary median
costae (read: parietal costae) mentioned by OehleRt

74 Paul SARTENAER
do not appear on the figures of the lectotype, formally
designated in the present paper, and on three specimens
from the Lower Devonian of Viré housed in the “Musée
des Sciences” of Laval.It is regrettable that subpareti,
which, according to OehleRt (1884, p. 415), is rather
frequently found in the Devonian outcrops of the
“Département de la Sarthe”, is poorly represented, and
more generally even not, in the available collections
from the Armorican Massif.
Proper collecting by regional geologists is needed
for improving our information on subpareti and the
Armorican pareti that are still insufficiently known and
not stratigraphically dated. It will then become possible
to deal with their generic status. In particular it will
allow finding out if the middle-sized R. subpareti from
the Laval Syncline belongs to the same genus as the
“nympha” of the same size and the larger “pareti” from
the Saint-Julien-de-Vouvantes Syncline. The problem
of the presence or absence of subpareti in Spain could
then also be solved.
An important conclusion of the present study is that
neither Inaequalibellirostrum pareti nor Stenorhynchia
nympha is present in the Armorican Massif.
Acknowledgments
This paper could not have been written without the help of many
colleagues. The author is grateful to Drs U. Jansen (Frankfurt
am Main) from the “Senckenberg Forschungsinstitut und Natur
Museum”, D. Korn from the “Museum für Naturkunde, Berlin”,
S. Long (London) from the British Museum (Natural History),
and J. Tréguier (Laval) from the “Musée des Sciences”, who
allowed him to browse among the collections under their care, and
to borrow some specimens. Dr. B. Wajsprych from the Institute
of Geological Sciences of the University of Wrocław kindly
made a cast of the lectotype (designated in the present paper)
of Terebratula Daleidensis figured by RoemeR, 1844. The late
Dr. H. Jaeger presented ten specimens of Oligoptycherhynchus
daleidensis to the author in June 1973 when the collections of the
“Preussische Königlich Geologische Landesanstalt” were in danger
of being destroyed. Dr. K.–W. Wenndorf (Braubach) generously
gave three specimens of Inaequalibellirostrum inauritum, and one
specimen of Oligoptycherhynchus hexatomus, from which serial
transverse sections could be made; he allowed these specimens to
be incorporated in the collections of the Belgian Royal Institute of
Natural Sciences, Brussels. Prof. F. Alvarez kindly offered a loan
of some specimens of Inaequalibellirostrum pareti. The author is
deeply grateful to Drs Ulrich Jansen and Klaus-Werner Wenndorf
for critically reading the typescript, and for their constructive
remarks.
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Paul SaRtenaeR
Department of Palaeontology
Royal Belgian Institute of Natural Sciences
Rue Vautier 29, B-1000 Brussels, Belgium
Typescript received: April 30, 2010
Revised typescript received: August 27, 2010

All figures are natural size
Explanation of Plate 1
Oligoptycherhynchus daleidensis (RoemeR, 1844)
Terebratula Daleidensis and related species
4
Figs 1-5 – Topotype A, IRScNBa12703. Dorsal, ventral, anterior, posterior, and lateral views. Costal formula: ; 0;
3
9
and
10
.
10 11
4 7 8
Figs 6-10 – Lectotype. Dorsal, ventral, anterior, posterior, and lateral views. Costal formula: ; 0; and . Dorsal,
3 8 9
anterior, and lateral views of this specimen have been figured by RoemeR (1844, pl. 1, figs 7a-c as Terebratula
Daleidensis).
Figs 11-15 – Topotype B, IRScNBa12704. Dorsal, ventral, anterior, posterior, and lateral views. Costal formula:
4
; 0;
9
.
3 10
Figs 16-20 – Topotype C, IRScNBa12705. Dorsal, ventral, anterior, posterior, and lateral views. Costal formula:
4
; 0;
7
.
3 8
Figs 21-25 –
4 9
Topotype D, IRScNBa12706. Dorsal, ventral, anterior, posterior, and lateral views. Costal formula: ; 0; .
3 10
Figs 26-30 –
4
Topotype E, IRScNBa12707. Dorsal, ventral, anterior, posterior, and lateral views. Costal formula: ; 0;
8
9
3 9
and .
10
Figs 31-35 – Topotype F, IRScNBa12708. Dorsal, ventral, anterior, posterior, and lateral views. Costal formula:
4
; 0;
9
.
3 10
83

84 Paul SARTENAER
plate 1